S,
BULLETIN OF
THE BRITISH MUSEUM
(NATURAL HISTORY)
GEOLOGY
VOL. XVII
1968-1969
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1970
DATES OF PUBLICATION OF THE PARTS
No. i. igth November . . . . . 1968
No. 2. igth November ..... 1968
No. 3. 1 7th January . . , . . 1969
No. 4. ryth January . . . . . 1969
No. 5. aist January ..... 1969
No. 6. 7th March ...... 1969
No. 7. I5th April ...... 1969
No. 8. i6th May ...... 1969
PRINTED IN GREAT BRITAIN
BY ALDEN & MOWBRAY LTD
AT THE ALDEN PRESS, OXFORD
CONTENTS
GEOLOGY VOLUME XVII
No. i. On the Cretaceous age of the so-called Jurassic Cheilostomatus Polyzoa
(Bryozoa). A contribution to the knowledge of the polyzoa-fauna of
the Maastrichtian in the Cotentin (Manche). E. VOIGT I
No. 2. The caudal skeleton in Mesozoic Acanthopterygian fishes. C.
PATTERSON 47
No. 3. Non-calcareous microplankton from the Cenomanian of England,
Northern France and North America. Part I : R. J. DAVEY 103
No. 4. A redescription of W. Carruthers ' type Graptolites. I. STRACHAN 181
No. 5. A revision of the English Wealden Flora, I Charales-Ginkgoales.
J. WATSON 207
No. 6. Two new Dicynodonts from the Triassic Ntawere Formation, Zambia.
C. B. Cox 255
No. 7. Lower Cambrian Archaeocyatha from the Ajax Mine, Beltana, South
Australia. F. DEBRENNE 295
No. 8. Some Bathonian Ostracoda of England with a revision of the Jones
1884, and Jones & Sherborn 1888 Collections. R. H. BATE 377
Index to Volume XVII 439
ON THE CRETACEOUS AGE OF
THE SO-CALLED JURASSIC
CHEILOSTOMATUS POLYZOA
(BRYOZOA)
A CONTRIBUTION TO THE KNOWLEDGE OF
THE POLYZOA-FAUNA OF THE MAASTRICHTIAN
IN THE COTENTIN (MANCHE)
E. VOIGT
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. i
LONDON: 1968
ON THE CRETACEOUS AGE OF THE SO-CALLER '
JURASSIC CHEILOSTOMATOUS POLYZOA
(BRYOZOA)
A CONTRIBUTION TO THE KNOWLEDGE OF THE
POLYZOA-FAUNA OF THE MAASTRICHTIAN
IN THE COTENTIN (MANCHE)
BY
EHRHARD VOIGT
Pp. 1-45; 8 Plates; 2 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. i
LONDON: 1968
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 17, No. I of the Geological
Palaeontological series. The abbreviated titles of the
Periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation
Bull. Br. Mus. nat. Hist. (Geol.).
© Trustees of the British Museum (Natural History) 1968
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 19 November 1968 Price £2
ON THE CRETACEOUS AGE OF THE SO-CALLED
JURASSIC CHEILOSTOMATOUS POLYZOA
(BRYOZOA)
A CONTRIBUTION TO THE KNOWLEDGE OF THE
POLYZOA-FAUNA OF THE MAASTRICHTIAN
IN THE COTENTIN (MANCHE)
By EHRHARD VOIGT
SYNOPSIS
The supposed Jurassic cheilostomatous Polyzoa described by J. W. Gregory (1894) as Mem-
branipora jurassica and Onychocella bathonica from the Bathonian of Ranville (Calvados), are
of Cretaceous age and must have come from the Maastrichtian of the Cotentin (Manche, France).
The matrices of the type specimens yielded thirty-three further species of Maastrichtian Polyzoa
of which nine are Cyclostomata and twenty-four Cheilostomata. Three new species Radulopora
minor n. sp., Rosseliana thomasi n. sp. and Frurionella fertilis n. sp., and the new genus Radulo-
pora are described.
CONTENTS
I. INTRODUCTION AND ACKNOWLEDGMENTS ..... 3
II. CHEILOSTOMATA OF CRETACEOUS AGE DESCRIBED FROM JURASSIC
BEDS .......... 4
III. POLYZOA FROM JURASSIC BEDS WHICH HAVE BEEN ERRONEOUSLY
REFERRED TO THE CHEILOSTOMATA ...... y
IV. ON THE ORIGIN OF GREGORY'S " JURASSIC " CHEILOSTOMATA . . IO
V. PALAEOZOIC CHEILOSTOMATA? . . . . . . . 13
VI. THE ACCOMPANYING FAUNA OF THE ROCK-MATRIX OF GREGORY'S
" JURASSIC " CHEILOSTOMATA . . . . . . . 14
VII. REFERENCES .......... 42
I. INTRODUCTION AND ACKNOWLEDGMENTS
THE predominant orders of Polyzoa (Bryozoa) of the Cretaceous and Cainozoic
periods are the Cyclostomata and the Cheilostomata. The Cheilostomata have been
increasing ever since an explosive development in the Upper Cretaceous, whereas
the Cyclostomata have decreased during the Tertiary and Quaternary following a
flourishing period in the Cretaceous. Today there exists only a comparatively
small relic of this cyclostomatous fauna stem which, in Jurassic times, was the
only living group of Polyzoa apart from some rare species of boring Ctenostomata.
This last statement contradicts all textbooks of palaeontology in which the
Cheilostomata are being erroneously recorded as beginning at the latest in the
Jurassic, an opinion repeated by R. S. Bassler (1953).
It is the purpose of this paper to prove that the so-called Jurassic Cheilostomata
described by Gregory (1894) from the Bathonian of Ranville (Calvados), are in fact
GEOL. 17, I. I
4 UPPER CRETACEOUS POLYZOA FROM COTENTIN
Upper Cretaceous in age and must have their origin in the Maastrichtian of the
Cotentin (Manche) in Normandy.
Other Jurassic Polyzoa described as Cheilostomata by several authors do not
belong to that order but are true Cyclostomata. In any case it must be emphasized
that, if they are undoubtedly Jurassic, they are not Cheilostomata, or if they belong
to that order they are not Jurassic.
I wish to record my warm appreciation of the late Dr. Dighton Thomas of the
British Museum (Natural History) to whom I am very much indebted for studies
in the Museum collection, for the loan of specimens, for discussion, and for his help
and advice. At the time of his death he had begun tidying the English of the
manuscript, and this was kindly finished by Dr. J. M. Hancock. I am also grateful
to Dr. A. B. Hastings for interesting discussions about the subject. Grateful thanks
are due to Prof. J. Lehman, Dr. J. Sornay and Dr. E. Buge of the Musee d'Histoire
Naturelle (Paris) for permission to study and to photograph the types of Bryozoa
in the d'Orbigny-Collection. I also wish to thank Dr. J. Roger, Paris, for kindly
accompanying me and guiding me in the Maastrichtian territory of the Cotentin
and other French classic localities. Finally my special thanks are due to the
Deutsche Forschungsgemeinschaft for financial help to further this research and
their support for my studies on Cretaceous Polyzoa.
II. CHEILOSTOMATA OF CRETACEOUS AGE DESCRIBED FROM
JURASSIC BEDS
Considering first the ages of undoubted Cheilostomata which have been described
as Jurassic forms, we can enumerate the following three species: Flustra flabelliformis
Lamouroux 1821, Onychocella bathonica Gregory 1894, and Membranipora jurassica
Gregory 1894.
i. " Flustra " flabelliformis Lamouroux 1821
1821 Flustra flabelliformis Lamouroux: 113, pi. 76, figs. 11-13.
This species, described in a footnote and figured by Lamouroux, apparently from
Ranville, is a fanlike fragment of a bilaminar Onychocella. The very simple figure
shows pyriform zooecia with round orifices increasing in size to the periphery of the
frond. An avicularium, rounded distally, seems to be indicated. Below the broken
cryptocyst the rectangular or box-like pattern of the zooecia is visible. The original
diagnosis of Lamouroux is:
"Flustra en forme d'eventail, fossile, epaisse, a bords entiers, composee de
deux membranes, une superieure mince, un peu translucide, divisee en alveoles
profonds, a bords irreguliers avec un oscule rond dans le centre, qui communique
a une cellule en forme de carre long, tres regulier, avec des cloisons epaisses
et solides, les transversales alternant entre elles, les longitudinales se prolongeant
sans interruption de la base aux extremites; grandeur, 2 a 3 centimetres;
epaisseur, environ i millimetre."
UPPER CRETACEOUS POLYZOA FROM COTENTIN 5
Unfortunately there is no possibility of restudying the typespecimen because,
according to Sherborn, Lamouroux's collection has been lost. Nevertheless, there
is no doubt that the specimen is not from the Bathonian of Ranville as supposed
by Gregory, but must be regarded as a Cretaceous Onychocella, likely to have come
from one of the Maastrichtian localities of the Cotentin (Manche), as is demonstrated
for the two following species. Lamouroux himself gives as the origin only the
neighbourhood of Caen, which perhaps includes not only Ranville but also the
Cotentin.
Gregory (1896 : 214) in his Catalogue of the Jurassic Bryozoa in the British
Museum suppressed his specific name bathonica for an Onychocella, regarded by him
in 1894 as a new species from the Bathonian of Ranville, in favour of Lamouroux's
species because he believed that the two forms were identical.
As will be shown, Gregory's Onychocella bathonica is the well-known Maastrichtian
Onychocella piriformis Goldfuss 1826 first described from the " Maastrichter Tuff-
kreide " in the Netherlands. If Gregory was correct in identifying his Onychocella
bathonica with Flustra flabelliformis Lamouroux, the latter name has priority. On
the other hand Lamouroux's figure is rather unsatisfactory, as Gregory himself
stated, for a conclusive identification of Flustra flabelliformis Lamouroux with
Onychocella piriformis Goldfuss. There are so many similar species of Onychocella
in the Upper Cretaceous that it seems to be quite impossible to find out which species
was intended by Lamouroux under this name. Gregory notes that Lamouroux's
figure is so indefinite that Pictet gave a figure of a form, which he referred to this
species, which was really Diastopora lamellosa Michelin. It is probable that Gregory,
in identifying his Onychocella bathonica with Flustra flabelliformis, was influenced
by his supposition that they are of Bathonian age, and as Jurassic species of Onycho-
cella must be very scarce, he believed that they must be identical.
2. " Onychocella bathonica " Gregory 1894
( = Onychocella piriformis Goldfuss 1826)
(PL 5, figs. 11-12)
1826 Eschar a piriformis Goldfuss: 23, pi. 8, fig. 10.
1851 Eschara piriformis (Goldfuss) von Hagenow: 75, pi. 9, fig. 6 and pi. n, fig. 6.
1894 Onychocella bathonica Gregory : 63, fig. 2
1896 Onychocella flabelliformis (Lamouroux) ; Gregory : 214, fig. 22.
1930 Onychocella piriformis (Goldfuss); Voigt: 454, pi. 16, figs. 1-2.
This is the supposed Jurassic cheilostomate which was described and figured by
Gregory under this name in 1894 and, as stated above, later treated by him as a
synonym of " Flustra "flabelliformis Lamouroux. The first mention of a " Jurassic
Onychocella " was made by Gregory (1893 : 239). It is represented in the British
Museum Collection by two specimens, D.iSi (type) and 0.480, both from the
Tesson-Collection. As previously suggested by the present author (Voigt 1930 : 454),
these two specimens are not from the Bathonian " Calcaire a polypiers " as recorded
6 UPPER CRETACEOUS POLYZOA FROM COTENTIN
by Gregory, but both are Upper Cretaceous in age. This is proved firstly by a
Cheilostomatous Polyzoan, Stamenocella marginata (d'Orbigny) which is visible in
the matrix of specimen D.i8i (PL 3, fig. i) and which was overlooked by Gregory,
and secondly by a large fauna of other Maastrichtian Polyzoa which has been
isolated from the adherent matrix of the block D.iSi and which is described in
section VI. This study shows clearly that all these supposed Bathonian Polyzoa
are in fact of Upper Cretaceous age as already indicated for " Membraniporajurassica "
by Lang (1916 : 96, 97 and 1922 : 197-198) and Larwood (1962 : 223).
Onychocella bathonica Gregory is the same species as Onychocella piriformis (Gold-
fuss) 1826 from Maastricht. It is quite commonly found in various localities of the
French Cotentin (Manche). Although Gregory has discussed Goldfuss's species
and written that it has a lower zooecial aperture, while the avicularian aperture is
larger and the front wall occurs only above and not on both sides of this, a comparison
with a specimen from Chef du Pont indicates that they are synonyms (pi. 5, fig. n).
The supposed differences are not significant and they are not found when material
from Maastricht or from the Cotentin localities is used for comparison. Some of the
opesiae1 in the figured (pi. 5, fig. 12) British Museum specimen D.iSi are a little
smaller than those of the figured specimen from Chef du Pont (pi. 5, fig. n), but
there are variations within the same specimen. Gregory's figure is deceptive because
it shows a small quadrangular fragment with only five whole zooecia and one avicu-
larium; it does not correspond in size to his two originals. The Holotype of Onycho-
cella bathonica Gregory is a large bent unilaminar frond of nearly 3 cm. length (pi. 5,
fig. 12). It contains a dozen avicularia and shows, on some zooecia, the very small
characteristic endozooecial ovicells which are just visible as minute swellings at
the distal ends of the zooecia figured here (pi. 5, fig. 12). The other specimen
0.480 is a large unilaminar fragment of n x 8-5 cm. size with four avicularia.
In discussing the affinities of Onychocella flabelliformis (Lamouroux) Gregory
maintains that its nearest ally may be von Hagenow's Cellepora (Discopora) konincki-
ana (1851 : 95, pi. n, figs. 11-12) from Maastricht, a species which, with its straight
rows of avicularia and small zooecia, is very differently shaped. In discussing
Onychocella ( = Cellepora} koninckiana (1896 : 215) he distinguished the form
figured by von Hagenow in his fig. n as a new species — Onychocella hagenowi. He
assigned it to a separate species because he believed that it has larger elliptical
opesiae with the longer axis longitudinal, an entire lower margin of the opesia and
much larger avicularia. This example shows how dangerous it is to judge the
variability of species on the evidence of figures alone. Von Hagenow was quite
correct in considering the two forms as only one species because they can often be
observed in the same zoarium as confirmed by Voigt (1930 : 460).
The first known species of Onychocella are from the Cenomanian, and they are
small and of a lower level of evolution. Judging from this point of view it would
be very odd if the oldest species should have the largest zooecia of the genus, more
than i mm. long, and big avicularia 1-3-1 -4 mm. in length as these are otherwise
developed only at the acme of the Onychocellids in the late Upper Cretaceous.
1 I have followed the advice of Dr. Hastings in using " opesia ", plural " opesiae ", rather than
" opesium ", plural " opesia ".
UPPER CRETACEOUS POLYZOA FROM COTENTIN 7
3. Castanopora jur assica (Gregory) 1894
(PI. 7, figs. 4-7)
1894 M embranipora jur assica Gregory: 62, text-fig, i.
1896 Membranipora jurassica Gregory: 212, text-fig. 21, p. 213.
1916 Rhiniopora jurassica (Gregory) Lang: 96.
1922 Rhiniopora scabra Lang: 196, pi. 4, fig. 7, text-fig. 62.
1922 Rhiniopora jurassica (Gregory) Lang: 197.
1962 Castanopora jurassica (Gregory) Larwood: 223, pi. 17, figs. 3-5, text-figs. 108-109.
Holotype D . 180, large bilaminar fragment of damaged zoarium partly embedded
in matrix. Upper Maastrichtian, Cotentin, Manche, France (not Maastricht,
Netherlands) .
Lang recognized the cribrimorph nature of this species. It was overlooked by
Gregory that the cribrimorph structure of the frontal-shield was preserved quite
well in some zooecia of the type specimen D.iSo (Tesson Coll.) which was figured
by Larwood (1962 : 224, text-fig. 108, pi. 17, fig. 5). Although Gregory described
it correctly as " erect foliaceous, bilaminate ", Lang defined this form as " encrusting
unilaminar " as did Larwood (1962). Dr. Dighton Thomas and Dr. A. Hastings
were kind enough to confirm my first observation made in the Museum collection
that Gregory's type specimen is bilaminar and not encrusting.
Gregory, misled by the erroneous data of Tesson's label, recorded the species as
coming from the Bathonian of Calvados. Lang (1922 : 197) recognizing the Maas-
trichtian age, supposed that it originated from the Dutch locality Maastricht itself,
and recorded the distribution of this species as, " Senonian, Maastrichtian, Maastricht,
Limburg, Holland ", specimen 0.3313 being labelled " Maastricht, Old collection ".
Larwood (1962, pi. 17, fig. 3) followed Lang and gave a photograph of this specimen.
I am much indebted to Dr. Dighton Thomas for lending me 0.3313, for there is no
other record of Castanopora jurassica from Maastricht, and the specimen may have
come from the same locality as D.iSo. Dr. Dighton Thomas compared the
matrices of these two specimens: they are very similar in colour, grain size, and
in size of fossil debris, and they could have come from the same locality in the
Cotentin.
As will be shown in section IV of this paper, the type locality cannot be Maastricht
in Holland, but must be the same as that of the last species, a locality in the Cotentin.
The matrix of the type specimen D . 180 is a hard, Polyzoan-bearing, detrital lime-
stone, resembling very much the " Craie a Baculites " or " Craie a Thecidees " of
the Cotentin. Lang has already recognized some other Cheilostomata in the matrix
of this block. They are described and figured here under the name of Multicrescis
laxata d'Orbigny, Rosseliana thomasi n. sp. and Pliophloea sp. These species are
unknown from Maastricht itself, and combined with the results above on " Onycho-
cella bathonica" , there can be no doubt that the two species both come from the
Cotentin.
All this is now confirmed by the fact that I have found four fragments of " Casta-
nopora " jurassica (Gregory) in my material from Chef du Pont (Cotentin, Manche).
8 UPPER CRETACEOUS POLYZOA FROM COTENTIN
Dr. Dighton Thomas kindly compared them with the holotype, and he has no doubt
that they represent the same species. They agree in measurements, in number
of costae, and all other characteristics. All are bilaminar with the exception of a
young zoarium which is unilaminar and whose zooecia are only 0-09-1-00 mm.
long. In the others the very large zooecia are about 1-2-1-4 mm. long. There are
about twenty-six to thirty-four costae with eight lateral costal fusions as stated by
Larwood. PL 7, fig. 5 shows an instructive view with some zooecia, one of which
has preserved its undamaged cribrimorph front wall; the others are broken and
show the " membranimorph " habit of Gregory's figure. On the reverse side of
the specimen (PI. 7, fig. 4) are some zooecia with the characteristic ovicells of the
genus which were previously not known in full preservation. They are hyper-
stomial and globular, prominent, and overlapping the distal zooecium as presumed
by Larwood. They are perforated by isolated fine pores (invisible in my figure)
as in Castanopora guascoi Ubaghs from Maastricht.
It must be emphasized that Castanopora jurassica (Gregory) has never been
found near Maastricht, although I have studied the Polyzoa of the Maastrichtian
Tuffkreide for 40 years. Near Maastricht another allied bilaminar form is repre-
sented, Cnstanopora guascoi (Ubaghs) (1865 : 51, pi. 2, fig. 3), whose zooecia in some
cases reach a length of 1-5 mm. but which has fewer costae (fifteen to twenty-two).
Specimens like these were described by von Hagenow as Cellepora (Dermatopora)
faujasi (von Hagenow : 1851, p. 99, pi. 10, fig. 19). It is impossible to mistake
Castanopora guascoi for Castanopora jurassica (Gregory), but it is of interest that,
if the costae are broken down, this species gives the appearance of Membranipora
bipunctata (Goldfuss 1826 : 26, pi. 9, figs. 7a-b), traces of costae being no longer
visible. The Cotentin fauna contains further allied bilaminar and unilaminar
species which could be mistaken for Castanopora jurassica, but there is little room
to discuss them. Rhiniopora scabra Lang 1916 from Riigen is, according to Larwood,
a synonym of Castanopora jurassica Gregory, because it agrees in general with the
number of costae, and in having 8 lateral costal fusions and pelmatidia.
In a recent work (Voigt 1968 : 65) finished before the issue of the present paper,
the genus Rhiniopora Lang 1916, united with Castanopora by Larwood (1962), is
maintained at least as a subgenus of Castanopora.
SPECIMENS :
0.3313. Large fragment partly embedded in matrix, recorded by Lang (1922 :
197) and figured by Larwood (1962, pi. 17, fig. 3), labelled " Maastricht, Old col-
lection ", but certainly from a locality of the Cotentin-Maastrichtian, Manche,
France.
0.49724. Small bilaminar fragment with three ovicelled zooecia. Upper
Maastrichtian, Chef du Pont, Cotentin, Manche, France. Collection E. Voigt.
0.49725. Young unilaminar zoarium with some damaged zooecia. Horizon
and locality as above. Collection E. Voigt.
Some fragments. Horizon and locality as above. In Collection E. Voigt, Ham-
burg, Nr. 3585 and 3924.
UPPER CRETACEOUS POLYZOA FROM COTENTIX g
III. POLYZOA FROM JURASSIC BEDS WHICH HAVE BEEN
REFERRED TO THE CHEILOSTOM AT A
It is now evident that the above mentioned Cheilostomata are not Jurassic.
There are scattered records in the literature of Cheilostomatous Polyzoa whose
Jurassic age is undoubted. In all such cases suspicion rises that they are not
Cheilostomata. The fallibility of many of these records depends on the progress
of science and today nobody can take seriously mention of Cheilostomatous genera
like Hippothoa, Eschara, Cdlaria or Cellepora etc. from the Jurassic by early authors.
However, Gregory (1894 : 61) in his note on the Jurassic Cheilostomata considered
Eschara ranvilliana Michelin from the Bathonian of Ranville, and Cellar ia smithi
Phillips from the Cornbrash of Scarborough, as true Cheilostomata and cited them
in support of his theory of the existence of Jurassic Cheilostomata. In his Catalogue
of the Jurassic Bryozoa (1896 : 56 and 127) Cellaria smithi Phillips is assigned to
Stomatopora and Eschara ranvilliana Michelin to Diastopora with no mention that
they were regarded as Cheilostomes by himself two years earlier.
F. D. Longe (1881) was still convinced that Eschara ranvilliana Michelin from the
Bathonian of Ranville belonged to the Cheilostomata when he wrote: " It is perfectly
clear, however, that some of the Oolitic Escharoids themselves possess the charac-
teristic cell features of the Cheilostomata in a marked degree; and their affinity to
the Cheilostomatous Escharidae has been recognized by no less authorities than
d'Orbigny and Michelin ".
He gave a detailed analysis of the supposed Cheilostomatous features of this
species and related forms, and tried to derive the Cheilostomata from certain Oolitic
Diastoporids whose zooecia remind one, by their oval or polygonal shape, of the
Cheilostomata. (" The decumbent cells in Diastopora may be regarded as ancestral
Cheilostomatous cells, and Diastopora itself as the parent stock from which many,
if not all, of the families of the Chalk and subsequent periods, grouped as Cheilo-
stomata, have been derived ".).
Walford (1894) published a note " On Cheilostomatous Bryozoa from the Middle
Lias ". Under the new generic name Cisternifera he described some species of cyclo-
stomatous Bryozoa with large heterozooecia — so-called " cistern-cells " — whose
relationship with the ovicells of the Cyclostomata was already assumed by Walford.
They were thought to bear, occasionally, minute avicularia on the upper lip of the
zooecia. Apart from the question of whether these structures are avicularia or not,
the different forms attributed to Cisternifera are true Cyclostomata, and Gregory
(1896) himself has distributed them amongst the genera Diastopora and Entalophora.
Cellepora davaiacensis Lissajous 1923 from the Bathonian of the Macon district
(France), from whose generic name one would expect it to be a Cheilostomatous
Polyzoan, and which Lissajous has compared with the Cheilostome Cellepora poly-
thele Quenstedt, has been recognized by David (1952) as belonging to the Calcispongia,
probably of the genus Synopelta Zittel. The list of literature on Jurassic Polyzoa
published by David (1960) contains many references to records of Cheilostomata in
other works.
From these investigations it is now certain that all supposed species of Jurassic
Cheilostomata prove to be mistaken identifications or erroneous stratigraphic records.
10
UPPER CRETACEOUS POLYZOA FROM COTENTIN
IV. ON THE ORIGIN OF GREGORY'S " JURASSIC CHEILOSTOM AT A '
In the Cotentin (Manche) Upper Maastrichtian a lithology similar to the " Tuff-
kreide " from Maastricht itself has long been known. The fades in the Cotentin
is a " tuffaceous " detrital limestone with remains of many Foraminifera, Polyzoa,
Echinoderms, Brachiopods (Craie a Thecidees ") or hard limestones (Craie a Bacu-
lites). It is true that certain blocks of these rocks can be easily mistaken for genuine
" Maastrichter Tuffkreide ", because many of the small fossils which make up the
rock are common to both strata.
An important difference is the absence of any larger foraminifera like Orbitoides,
Lepidorbitoides , Siderolites or Omphalocyclus etc. which are distinctive of the Upper
Maastrichtian in Holland and Belgium. Hofker (1959) in his monograph on the
Foraminifera of the Cotentin Maastrichtian has shown that these beds must be
intermediate in age between horizons Cr 4 and Mb in the terminology of Uhlenbroeck
for the Upper Cretaceous in South Limburg. This would indicate a stratigraphic
position between the phosphatic chalk of Ciply and the base of the Tuffaceous chalk
of St. Symphorien in the Mons basin, or an equivalent of the higher beds of Folx-les-
Caves and Orp-le-Petit in northern Belgium. This could explain the lack of larger
Foraminifera in this region which did not invade the northern regions before the
higher horizons of the Maastrichtian. The Maastrichtian age is based upon the
occurrence of Scaphites constrictus J. Sowerby in the area of Valogne. For the other
fauna see Vieillard & Dollfus 1875.
In 1957 I visited the Cotentin region in order to study the classic Senonian localities
of d'Orbigny (1850-54) who had described from here, chiefly from Nehou and from
Sainte Colombe, some hundred species of Polyzoa. Although there were no extant
exposures in the immediate neighbourhood of these villages, some larger quarries
near Fresville and Port Filiolet and an excavation for the dairy of Chef du Pont
supplied material which furnished examples of a considerable part of the Maas-
trichtian Polyzoa which were described and figured by d'Orbigny. (For the site
Valognes
Nthou
zter
Fresville
* ^&
S* Colombe
Rauvilte -
la Place °*j
Rue-Tourville
24 6 d 10
)
Chef du Pont
Port Filiolet
km
FIG. i. Map of the outcrops (black) and the localities of the Maastrichtian in the Cotentin
area (Normandy). (After Carte geologique detaillee de la France (i : 80000) Feuille 28,
St. L6 (2 erne Ed.) 1926.
UPPER CRETACEOUS POLYZOA FROM COTENTIN n
of these localities see Text-fig, i.) This material was augmented by some samples
given by Dr. F. Schmid (Hannover) and Polyzoa from Fresville presented by Dr.
P. Marie (Paris). It is interesting that many different and new forms not mentioned
by d'Orbigny were found at these localities, and by contrast, a large number of
d'Orbigny's species could not be found again. Perhaps there are some horizons with
a different fauna, and the beds of Sainte Colombe and Nehou may belong to such
levels, whose exposures are abandoned today, or the different localities represent
heterogeneous ecologic biotopes.
The Polyzoan fauna of this region is the most important one in the northern area
and its knowledge is fundamental to our knowledge of the Maastrichtian Polyzoa.
With the exception of the inadequate revision of the Cretaceous Polyzoa of d'Orbigny
as a whole, which was undertaken by Pergens (1889) and by Canu (1900), nobody
has concerned himself with the Polyzoan fauna of the Cotentin for no years. An
up-to-date revision of this fauna is therefore an urgent task, but it cannot be under-
taken without a re-study of d'Orbigny's types which are preserved in the Musee
d'Histoire naturelle in Paris.
The most obvious obstacle for a successful revision is the fact that many of
d'Orbigny's type specimens are missing, or, if specimens are present, it is often
impossible to say with certainty whether they are true " types " or not. On the
other hand many of the drawings do not agree well with the originals ; they may be
stylized, restored or improved and consequently it is often rather difficult to give a
satisfactory identification of d'Orbigny's species (see Canu 1900 : 335). His omission
to specify the exact locality of the figured specimen in cases when there is more than
one such locality, increases the difficulty of identifying the types. There are about
300 photographs of d'Orbigny's polyzoan types of the Paleontologie Fran9aise in
the author's collection, and this stock, together with a rich collection of French
Cretaceous Polyzoa, are the basis for the following discussion.
The proofs for a Cotentin origin for Gregory's " Jurassic Cheilostomata " are:
1. The matrix of sediment adhering to Gregory's originals is identical to that of
the above-listed Maastrichtian localities of Cotentin. The matrix detached from
block D.i8i with Onychocella piriformis Goldfuss has yielded two specimens
(66.42981-82) of the characteristic fossil Thecidea papillata V. Schlottheim (pi. i,
figs. 9-10) and it is evident that it has been collected from the so-called " Craie a
Thecidees " of the Cotentin.
2. The objection that this brachiopod is also common at Maastricht and in Belgium
in a similar facies is weakened by the fact that the accompanying fauna in this
block, consisting of thirty-one species of Polyzoa, contains some species which are
confined to the Cotentin fauna and have never been found in Holland and Belgium,
although indeed both strata have many species in common. Examples of species
not known from these countries are: Multicrescis laxata d'Orbigny, Membranipora
unipora Marsson, Stamenocella cf. marginata (d'Orbigny), Castanopora jurassica
(Gregory), Onychocella bellona (d'Orbigny), Onychocella cf. cepha d'Orbigny, Onycho-
cellaria caecilia sp., Semiescharinella complanata d'Orbigny, Lunulites sp., Rosseliana
thomasi n. sp., Frurionella fertilis n. sp. and others.
12 UPPER CRETACEOUS POLYZOA FROM COTENTIN
3. A Cotentin origin is further suggested by another Cretaceous Polyzoan in the
museum collection which is embedded in a typical sample of " Craie a Thecidees ".
It is a large specimen of a Reticrisina, determined as the Jurassic " Reticulipora
dianthus (Blainville) " (6.4569 Old collection) and labelled " Bathonian Fresville ".
This example gives a good idea of how the error could have arisen : Bathonian rocks
being absent near Fresville, Cretaceous Polyzoa were mistaken for Bathonian fossils.
Gregory's " Jurassic Cheilostomata " both belong to the Tesson-Collection, which
according to Gregory's Catalogue contains many Bathonian Polyzoa from the famous
locality of Ranville (Calvados).
The two localities are about 100 km. apart, and it is probable that the labels
were confused or that the Cretaceous Polyzoan limestone was mistaken for the not
dissimilar Coral and Polyzoan-limestone (" Calcaire a polypiers ") of the Bathonian.
The Tesson-Collection to which Gregory's types belong was acquired in 1857
(Gregory 1896 : 35) and the fossils were probably collected many years ago at a
time when geological mapping of this region was still in its infancy.
It is not impossible that Gregory's types both came from Fresville like the Reti-
crisina mentioned above. The rock matrix of D.iSi is very like that of the " Craie
a Thecidees " of Fresville, but as Dr. Dighton Thomas informed me, that from Chef
du Pont is even closer in appearance. But such lithological differences may occur
in the same sequence, and are of no great importance.
At least one other possible explanation of this error should be mentioned. Between
the Maastrichtian outcrops of Sainte Colombe and Rue Tourville is the village
Rauville-la-Place (see Text-fig, i). Possibly a label " Rauville " was misunderstood
for " Ranville ", Calvados.
It must be reckoned that errors like these are more common in old collections, and
it may be recalled that a considerable number of Reuss' " Cenomanian Polyzoa of
Saxony" came from the Vincentown Limesand in New Jersey (U.S.A.), of Dano-
Paleocene age (Voigt 1942). Another mistake is the supposedly new " Polyphyma
bulbosa " Hamm (1881) from Maastricht which is a Hauterivian form from northern
Germany, and which was described in 1839 by Roemer as Alveolites heteropora
(Voigt 1953 : 57). Certain suspicious anachronisms of phylogenetic level may be
explained in this manner as has been shown by the present author with some of
Reuss' types.
4. I have collected much material of Bathonian Polyzoa at the classic locality of
Ranville (Calvados), but I have never found any trace of Cheilostomata. The
objection, that the Cheilostomata may originate in Jurassic times cannot be con-
tested; but we have no remains of them earlier than the Lower Cretaceous. It is
true that in the last decade several important groups of fossils have been proved to
be older than was formerly assumed. Today we know of Cambrian bivalves,
Carboniferous belemnites, Triassic frogs, and these examples could be augmented.
It is possible that Jurassic Cheilostomata may be found one day. But in such a
case we should expect primitive forms like Membrammorphs as are found in the
lower Cretaceous and not highly developed forms like Onychocella, or highly special-
ized forms like the Cribrimorph Castanopora.
Lower Cretaceous Cheilostomata are of the greatest phylogenetic interest, and all
UPPER CRETACEOUS POLYZOA FROM COTENTIN 13
forms described to date are rare. Most of them are Albian and the oldest is recorded
from the Neocomian. Their number is so small that they can be listed in a few lines :
1. Membranipora neocomiensis d'Orbigny 1853,
Neocomian, Saint-Sauveur, Yonne, France. The only specimen was too bad
to be figured by d'Orbigny. It must be regarded as a nomen nudum.
2. "Membranipora" constricta d'Orbigny 1853,
Aptian, Les Croutes (Yonne).
Remarks: I have photographed the supposed " type " of this species (pi. 7,
fig. 8). It is without doubt a " Membranipora " s.L, but it does not cor-
respond with the original figure.
3. Rhammatopora (?) johnstoniana Mantell 1844,
Aptian, Lower Greensand, Shanklin-Sand, Kent.
4. Rhammatopora gaultina Vine 1890, with his synonyms Rhammatopora vinei
Lang and Rhammatopora pembrokiae Lang (c.f. Thomas & Larwood 1960)
Albian-Cenomanian, England.
5. Charixa vennensis Lang 1915, Albian, Dorset.
6. Pyripora texana Thomas & Larwood 1956, Albian, Texas (U.S.A.).
7. Wilbertopora mutabilis Cheetham 1954, Albian, Texas (U.S.A.).
Vine (1890) mentioned Membranipora fragilis d'Orbigny, Membranipora ? obliqua
d'Orbigny, Membranipora elliptica v. Hagenow and Hippothoa simplex d'Orbigny
from the Red Chalk of Hunstanton. The identification of these forms must be
revised, but nevertheless all the named species are primitive encrusting membrani-
morphs of the Division Malacostega, suborder Anasca, and half of them are mono-
or oligoserial, and lack either ovicells or avicularia as we must theoretically presume
if our ideas about the evolution of the early Cheilostomata are correct. They are
followed in the Cenomanian by the first primitive Cribrimorphs and Coilostega with
the families of Onychocellidae and Microporidae. Therefore it is hard to under-
stand why Canu & Bassler (1920 : 318) stated that following the Membraniporae,
the Acroporidae Canu 1913 (= Porinidae d'Orbigny 1852) of the suborder Asco-
phora are the most ancient Cheilostome fossils. The main evolution of this family
is in the later Upper Cretaceous, and the only described species of Porina from the
Cenomanian is P. cenomana Lecointre (1912) whose origin and inner structure needs
revising.
V. PALAEOZOIC CHEILOSTOMATA?
In this connection the question of the systematic position of the north American
families Worthenoporidae Ulrich 1893 (Carboniferous) and Palescharidae Miller 1889
(Ordovician-Devonian) cannot be neglected. These were established for the single
genera Worthenopora Ulrich 1889 and Paleschara Hall 1874 which resemble Cheilo-
stome Polyzoa in some features. Ulrich (1890) stated that the affinities of that genus
are nearer to the Membraniporidae than any other and that his present views
would admit it being placed in the Cheilostomata. Nickles & Bassler (1900) regarded
both families without any restriction as Cheilostomata. This classification was
followed by many authors, but in recent times this opinion seems to have been
I4 UPPER CRETACEOUS POLYZOA FROM COTENTIN
abandoned. Bassler (1953) has placed them among the Cryptostomata, but con-
siders Worthenopora " may belong among cheilostomes ".
Worthenopora, indeed, has the outward aspect of a Cheilostome with its triangular
or semielliptical apertures, with posterior raised margin and spine bases. Paleschara
with its simple short polygonal zooecia is like a Membranipora which possesses
completely opened opesiae without any trace of a gymnocyst.
Dr. Dighton Thomas has kindly lent me some specimens of both genera from the
museum collection. Although there is no space here to go into details, further studies
of this problem are intended. I have got the impression that they cannot be attri-
buted to the Cheilostomata. Paleschara must be regarded as a very primitive
Cryptostome, and Worthenopora, in contrast, as a specialized one. The latter has
reached a level of evolution which reminds one of certain Cheilostomata. It has
not been found in beds younger than Mississippian, and there are no intermediate
forms between it and the Cretaceous Cheilostomata. Consequently it seems to be
impossible to regard it as an ancestor of the true earliest membranimorph Cheilo-
stomata of the Lower Cretaceous.
Paleschara, with its network of rather simple polygonal zooecia, may perhaps be
primitive enough to give rise to Cheilostomata-like forms. But it could be better
regarded as a cryptostomatous form corresponding to a membranimorph level of
primitive Cheilostomata.
We have no palaeontological evidence for a descent of the Cheilostomata from the
Cryptostomata or from the Ctenostomata. As Borg (1930 : 54) and Cori (1941)
regard the Ctenostomata as emanating from primitive Cheilostomata, it must be
emphasized that the Ctenostomata are the older group, represented by fossils from
the Ordovician onwards and that there is no possibility of deriving the Ctenostomata
from the Cheilostomata. Silen (1942) regards the Cheilostomata and the Cteno-
stomata as closely allied (" Cheilo-Ctenostomata ") and believes that both have
common ancestors.
Silen has established an interesting theory about the origin of the Cheilostomata
from hypothetical primitive forms like the recent Labiostomella which he calls
Protocheilostomata. These have erect zoaria with frontal budding and other
primitive features, and it would mean that the encrusting growth of many Cheilo-
stomata and the lateral budding as existing in all other Cheilostomata, are secondary.
It should be emphasized that the oldest known Cheilostomata from the Lower
Cretaceous do not show this primitive character. All are encrusting, and they
must already have passed the evolutionary level of the " Protocheilostomata ",
which according to Silen were feebly or not at all calcified. If this is admitted,
there remains no possibility of regarding Worthenopora or Paleschara as early Cheilo-
stomata of the Palaeozoic.
VI. THE ACCOMPANYING FAUNA OF THE MATRIX OF GREGORY'S
" JURASSIC CHEILOSTOMATA "
It is satisfactory that in the matrix of Gregory's " Jurassic Cheilostomata " are
enclosed other Bryozoa which were overlooked by Gregory. In block D.iSo,
containing Castanopora jurassica, Multicrescis laxata d'Orbigny, Rosseliana thomasi
n. sp. and Pliophoea sp. could also be observed.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 15
I am much indebted to Dr. Dighton Thomas for his permission to remove about
2 cm.3 of the matrix of block D.iSi with Onychocella bathonica. This piece was
cracked under a press and the washing of the residue has yielded a fauna of thirty-
three species of Polyzoa and one Brachiopod. This fauna is described in detail in
the following part of this paper. Most of the treated specimens are figured although
the state of preservation is rather poor because the hard rock is unfavourable for
clean preparation. All specimens are somewhat damaged, or rolled and, unfortu-
nately, recrystallized or covered with minute calcite crystals as is often observed in
the Cotentin material, and it is nearly impossible to stain them with colour. This
explains any mediocre photos. In many cases figures of comparable specimens of
the same species from the Cotentin are given for comparison. If they do not always
present exactly the same picture as the specimens from the original French localities,
this is because there often exists a great variability in size or growth-stage, and
in preservation, and that the identification is based upon a vast quantity of material.
It is surprising that in only 2 cm.3 of rock from D . 181 some small fragments of new
species were found, although these have long been known to the author from the
Maastrichtian of the Cotentin. The opportunity is taken here to describe them.
But it should not be forgotten that this small fauna from only a few cm.3 matrix
must represent a very small part of the rich Polyzoan fauna of the Cotentin Maas-
trichtian, and that its composition is purely accidental.
The list given in table i, p. 41, contains thirty-three Polyzoan species of which
twenty-four are Cheilostomata. It is not complete because in some cases a com-
plete identification could not be made, and for a few forms no identification was
possible. Excepting Pliophloea sp. all forms have been found by the author in
the Maastrichtian of the Cotentin. Twenty-six species were found at Chef du
Pont, eighteen at Port Filiolet and nine at Fresville. But it should be taken into
account that the investigated material is too poor for important deductions. Many
species very common at all Cotentin localities are not represented, and others are
new for the Cotentin. The affinities with the classic locality of Maastricht are
proved by fourteen species but this figure may be too large because several forms of
both regions may be identical. Von Hagenow's types from Maastricht having been
lost during the second world war, it is not yet possible to decide this question with
certainty.
I. POLYZOA
A. CYCLOSTOMATA
Genus BERENICEA Lamouroux 1821
i. Berenicea sp.
One small incomplete zoarium which is indeterminable, encrusts Reteporidea
lichenoides Goldfuss.
SPECIMENS. A minute incomplete zoarium encrusting Reteporidea lichenoides
Goldfuss (=0.49569).
16 UPPER CRETACEOUS POLYZOA FROM COTENTIN
Genus ENTALOPHORA Lamouroux 1821
2. Entalophora benedeniana (von Hagenow)
1851 Pustulopora benedeniana von Hagenow: 17, pi. i, fig. 6.
1899 Entalophora madreporacea Goldfuss var. benedeni von Hagenow; Gregory: 239.
1964 Entalophora benedeniana (von Hagenow) Voigt: 422, pi. i, figs. 1-7.
One small fragment embedded in matrix has been referred to this well-known
species from Maastricht which is represented in the author's collection from the
Maastrichtian of Chef du Pont (Manche). For morphological details and ovicells
see Voigt (1964).
SPECIMENS. 0.49560. One poorly preserved fragment in matrix from D.iSi.
Upper Maastrichtian, Cotentin, Manche, France.
Genus IDMIDRONEA Canu & Bassler 1920
3. Idmidronea macitenta (von Hagenow)
1851 Idmonea macilenta von Hagenow: 29, pi. 2, fig. 4.
?i85i Idmonea ramosa d'Orbigny: 736, pi. 6n, figs. 11-15.
1899 Retecava ramosa (d'Orbigny) Gregory: 192, pro parte.
1951 Idmidronea macilenta Voigt: 38, pi. 4, figs. 14-17.
One small distal fragment of this abundant species of the Upper Maastrichtian
belongs to /. macilenta. These thin distal branches do not have the numerous
firmatopores which are well developed on the reverse side of the broader and older
stems. Therefore they are very similar to Idmonea (Tubigera) antiqua Defrance
(d'Orbigny 1853 : 722, pi. 613, figs. 11-15, figured under the name of Stichopora
regularis d'Orbigny). Idmonea disticha Goldfuss (sensu von Hagenow 1851 : 30,
pi. 2, fig. 8) may be mistaken for this form also, but always it is not so flat as the
distal ends of Idmidronea macilenta. The species cited by d'Orbigny from Sainte-
Colombe under the latter name may be Idmonea macilenta.
If this species is con-specific with Idmonea ramosa d'Orbigny, the latter name has
priority and must be preferred. It is represented at Fresville (Cotentin, Manche).
STRATIGRAPHICAL RANGE. Upper Maastrichtian.
SPECIMENS. 0.49561. A worn fragment from matrix of D.iSr. Upper
Maastrichtian, Cotentin, Manche, France.
0.49843. A fragment, upper Maastrichtian, Chef du Pont, Cotentin, Manche,
France. E. Voigt Collection.
Genus HETEROCRISINA Gabb & Horn 1860, em. Voigt 1964
4. Heterocrisina communis (d'Orbigny)
1853 Idmonea communis d'Orbigny: 745, pi. 750, figs. 6— 10.
1887 Idmonea pseudodisticha (non von Hagenow) Marsson: 28, pi. 2, fig. 8.
1860 Heterocrisina abbottii Gabb & Horn: 404, pi. 69, figs. 45-47.
1899 Retecava abbottii (Gabb & Horn) Gregory: 205.
non 1907 Idmonea abbottii (Gabb & Horn) Ulrich & Bassler: 321, pi. 22, figs. 3-4.
1964 Heterocrisina communis (d'Orbigny) Voigt: 432 pi. 3, figs. i-io.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 17
Two small worn fragments belong to this species, which is very distinctive because
of its large frontal ovicell, although it has often been mistaken for other species.
For further information about its generic position and morphological details see
Voigt (1964).
This form is common at all Maastrichtian localities in the Cotentin (Fresville,
Port Filiolet and Chef du Pont).
STRATIGRAPHICAL RANGE. Campanian-Maastrichtian.
SPECIMENS. D . 49562-63. From matrix of D . 181. Upper Maastrichtian, Coten-
tin, Manche, France.
Genus CRISISINA d'Orbigny 1847
5. Crisisina carinata (Roemer)
(PL i, figs. 4-5)
1840 Idmonea carinata Roemer: 21, pi. 5, fig. 20.
1964 Crisisina carinata (Roemer) Voigt: 429, pi. 4, figs. 1-7. (See full references.)
Three worn fragments belong to this very common and widespread species of the
Upper Cretaceous. The synonymy is very confused — see Voigt (1964) — where all
known synonymies are given and the generic classification is discussed.
Very abundant at all Maastrichtian Cotentin localities.
STRATIGRAPHICAL RANGE. Cenomanian-Paleocene.
SPECIMENS. 0.49564-66. From matrix of D.iSr. Upper Maastrichtian, Coten-
tin, France.
0.49567. A fragment for comparison with 0.49566. Upper Maastrichtian,
Port Filiolet, Cotentin, Manche, France, Voigt Collection.
Genus OSCULIPORA d'Orbigny 1847
6. Osculipora truncata (Goldfuss)
1826 Retepora truncata Goldfuss: 28, pi. 9, fig. 14.
1851 Truncatula truncata (Goldfuss) v. Hagenow : 35, pi. 3, fig. 2.
1851 Truncatula tetrasticha von Hagenow: 34, pi. 3, fig. 3.
1909 Osculipora truncata (Goldfuss) Gregory: 58.
1922 Osculipora truncata (Goldfuss) Canu & Bassler: 57, pi. 23, figs. 1-6.
One small worn fragment may be referred to 0. truncata; it is in the condition of
Truncatula tetrasticha von Hagenow, which represents highly worn branches of
0. truncata (Goldfuss.)
Further investigations are necessary to check whether the older citations of this
form from the Cenomanian by Reuss (1872) are correct. In the author's collection
from Chef du Pont, Port Filiolet and Fresville are numerous fragments of this
species, which was not recorded by d'Orbigny from any of his Cotentin localities.
STRATIGRAPHICAL RANGE. Cenomanian (?) to Maastrichtian.
SPECIMENS. 0.49568. A very poorly preserved worn fragment from the matrix
of D.iSi. Upper Maastrichtian, Cotentin, Manche, France.
GEOL. 17, I. 2
i8 UPPER CRETACEOUS POLYZOA FROM COTENTIN
Genus RETEPORIDEA d'Orbigny 1849
7. Reteporidea lichenoides (Goldfuss)
1826 Retepora lichenoides Goldfuss: 29, pi. 9, figs, isa-b.
1851 Idmonea lichenoides (Goldfuss) von Hagenow: 28, pi. 2, fig. 6.
1899 Retecava lichenoides (Goldfuss) Gregory: 194, fig. 16, p. 195.
One very poor fragment which is encrusted by a small young Berenicea, has been
recognized as this species which is very abundant in the Upper Maastrichtian. It
is represented in the author's collection from Port Filiolet by some specimens. It
has not previously been recorded from the Cotentin Maastrichtian.
STRATIGRAPHICAL RANGE. Maastrichtian.
SPECIMENS. 0.49569. A small worn fragment with an encrusting Berenicea
from the matrix of D.iSi. Upper Maastrichtian, Cotentin, Manche, France.
Genus PETALOPORA Lonsdale 1850
8. Petalopora sp.
(PI. i, figs. 1-3)
There are a few fragments of a badly preserved Petalopora which are conspecific
with similar specimens from Chef du Pont (pi. i, fig. 2) . Their identification involves
some difficulties because it is impossible to identify them from published figures.
They are allied to Heteropora reiiculata Marsson (1887 : 26, pi. 2, fig. 4) in the size
of the branches and diameter of the apertures (about 0-14-0-15 mm.), but the
mesopores are much less conspicuous, and on the figured fragment (pi. i, fig. i)
they look a little like longitudinally-oriented lines which are straight or sinuously
bent and interspersed between fine ribs. On the specimen from Chef du Pont the
mesopores are larger and they show a more longitudinally-oriented inconspicuous
network (pi. i, fig. 3). I have no doubt that these three specimens are conspecific
in spite of this difference but I dare not identify them with any known species.
SPECIMENS. 0.49570-72. Three small worn fragments from the matrix of
D.iSi. Upper Maastrichtian, Cotentin, Manche, France.
0.49573. A larger fragment from the matrix of D.iSi. Upper Maastrichtian,
Cotentin, Manche, France.
0.49574. A larger fragment. Upper Maastrichtian, Port Filiolet, Cotentin,
Manche, France. Voigt Collection. One branched fragment. Upper Maastrich-
tian, Chef du Pont, Cotentin, Manche, France. Voigt Collection, Hamburg, Nr. 3969.
Genus MULTICRESCIS d'Orbigny 1854
9. Multicrescis laxata d'Orbigny
(PL i, figs. 6-8)
1854 Multicrescis laxata d'Orbigny: 1077, pi. 800, figs. 10-11.
HOLOTYPE. Upper Maastrichtian, Sainte Colombe, Cotentin, Manche, France.
d'Orbigny Collection, Paris, Musee d'Histoire Naturelle Nr. 8416.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 19
There is a fragment of a " heteroporid " Polyzoan with an encrusting colony of
Rosseliana thomasi n. sp. embedded in the matrix of Castanopora jurassica Gregory
(D.i8i). It cannot be distinguished from d'Orbigny's type specimen of Multi-
crescis laxata, whose apertures and the mesopores are exactly the same. Unfortu-
nately there are no other specimens of this species in d'Orbigny's collection which
would allow confirmation of the inner structures by sections. Specimens from
Chef du Pont (Manche), which seem to be conspecific with d'Orbigny's species have
a median lamella like Grammascosoecia Canu & Bassler 1922. Pergens (1889 : 373)
and Canu & Bassler (1922 : 119) have included this species in the synonymy of
Grammascosoecia dichotoma (Goldfuss) from Maastricht (see von Hagenow 1851 : 47,
pi. 5, fig. 15). This might be correct; some specimens from Fresville can hardly be
distinguished from the Maastricht species. On the other hand there are some fifty
fragments from Fresville, and none show the characteristic pattern of small regular
smooth quadrangles which grow from the calcified mesopores (cf. von Hagenow's
fig. I5i and Voigt 1951 pi. 4, fig. i) and which can be observed in the majority of
the Maastricht specimens. Therefore I still hesitate to unite it with Gramma-
scosoecia dichotoma (Goldfuss).
D'Orbigny attributed his species to his genus Multicrescis which is multilamellar.
But this is not the case in M. laxata, although d'Orbigny noted two layers in his
type-specimen from Sainte-Colombe, which is a basal fragment (pi. I, figs. 7-8).
It is well known that the basal stems often develop more than one layer of zooecia,
and therefore there is no reason to place this form in the genus Multicrescis. Many
specimens from Fresville have radiating rows of peristomes as in Multicavea, which
can also be observed in Grammascosoecia dichotoma (Goldfuss). But the ovicell of
the Cotentin form has not yet been discovered. Therefore I prefer to leave this
form provisionally under the name given by d'Orbigny.
It should be noted that the median lamella in Grammascosoecia dichotoma (Gold-
fuss) is not constant. It is lacking in many specimens from Maastricht, and there
is no reason to assume a different species.
STRATIGRAPHICAL RANGE. Upper Maastrichtian.
SPECIMENS. 0.49575. A worn fragment with encrusting Rosseliana thomasi
n. sp. embedded in matrix of Castanopora jurassica (Gregory) (D.iSo). Upper
Maastrichtian, Cotentin, Manche, France. Labelled erroneously by Gregory as
" Bathonian, Ranville, Calvados ".
B. CHEILOSTOMATA
MEMBRANIPORA Blainville 1830 (sensu lato)
10. " Membranipora " unipora (Marsson)
(PL 2, figs. 7-8)
1852 Flustrella simplex d'Orbigny: 293, pi. 699, figs. 14-16.
1887 Biflustra unipora Marsson: 52.
1929 Membranipora genucia Brydone: 37, pi. 13, figs. 10-11.
20 UPPER CRETACEOUS POLYZOA FROM COTENTIN
1930 Membranipora unipora (Marsson) : Voigt 420, pi. 4, fig. 5.
1925 Membranipora unipora (Marsson): Levinsen 329, pi. 2, fig. 22.
HOLOTYPE. Upper Maastrichtian, Nehou, Cotentin, Manche, France. d'Orbigny
Collection, Paris. Musee d'Histoire Naturelle Nr. 8130.
One unilaminar specimen embedded in matrix, showing some zooecia with the
characteristic avicularium on the gymnocyst below the opesia and the narrow
helmet-shaped hyperstomial ovicell, agrees very well with d'Orbigny's type from
Nehou (pi. 2, fig. 8) and another well preserved specimen from the Maastrichtian of
Port Filiolet (Manche). The zoarial length is from 0-80-1-20 mm. The spines of
the well raised margin of the opesia, which is 0-45-0-50 mm. long, number about
twenty, but they are inconspicuous and often hidden by recrystallization of calcite.
The zooecia and the opesiae of my specimen of this species from Port Filiolet are a
little larger than those of the Museum-specimen but this lies within the range of
variation of this form. Membranipora genucia, described by Brydone from the
upper Campanian of Meudon near Paris, is a synonym as shown by comparison
with Cotentin and Rugen specimens with those from Meudon.
Marsson who translated this species from Flustrella to Biflustra, has changed
the species name simplex to unipora because there existed already a recent Biflustra
simplex d'Orbigny 1839. Otherwise another Cretaceous Membranipora simplex
d'Orbigny exists too. It is clear that the placing of this species in " Membranipora "
is only provisional; a revision of the Cretaceous Membraniporae might put it in
another genus.
STRATIGRAPHICAL RANGE. Upper Campanian-Maastrichtian.
SPECIMENS. 0.49576. Fragment with ovicelled zooecia from the matrix of
D.iSr. Upper Maastrichtian, Cotentin, Manche, France.
Genus AMPHIBLESTRELLA Prud'homme 1960
ii. Amphiblestrella elegans (von Hagenow)
(PL 4, figs. 1-3)
1851 Siphonella elegans von Hagenow: 84, pi. 6, fig. 7.
1851 Flustrella baculina d'Orbigny: 291, pi. 699, figs. 4-6.
1930 Amphiblestrum elegans (von Hagenow) Voigt: 448, pi. 13, figs. 13-16.
1960 Amphiblestrella elegans (von Hagenow): Prud'homme: 949.
1962 Amphiblestrum elegans (von Hagenow) Berthelsen: 100, pi. 9, figs. 1-5.
There is only one small fragment which is 1-7 mm. long and shows eight rows
of zooecia; normally there are ten to sixteen. The zoarial dimensions are smaller
than those of specimens from the type-locality of Maastricht, and from the Cotentin,
where this species is very common. The axial canal of the hollow zoaria, clearly
visible in the thicker zoaria, is much reduced in the slender branches, and may disap-
pear almost completely as shown in d'Orbigny's type specimen of his Flustrella
baculina (pi. 4, fig. 2) from Nehou. The zooecia from Danian material are mostly
longer than those from the Maastrichtian (about 0-7 mm. instead of 0-6 mm.). The
size of the opesiae is very variable in this species. Canu (1900) in his revision of
UPPER CRETACEOUS POLYZOA FROM COTENTIN 21
d'Orbigny (1851-54) has incorrectly regarded this species as a synonym of Flustrella
irregularis d'Orbigny.
In the Cotentin localities this species is represented from Fresville and Chef du
Pont and by d'Orbigny's type specimen of Flustrella baculina from Nehou.
STRATI GRAPHICAL RANGE. Maastrichtian-Danian.
SPECIMENS. D. 49577. A worn fragment from matrix of D.iSi. Upper
Maastrichtian, Cotentin, Manche, France.
D. 49578. A well preserved branched fragment. Upper Maastrichtian, Chef du
Pont, Cotentin, Manche, France. Voigt Collection.
Type-specimen of Flustrella baculina d'Orbigny. Upper Maastrichtian Nehou,
Cotentin, Manche, France. In d'Orbigny Collection, Paris, Musee d'Histoire
Naturelle Nr. 8127.
Genus RADULOPORA nov.
DERIVATIO NOMINIS. Derived from the species-name of Biflustra radula Marsson
1887.
DIAGNOSIS. Zoarium bilaminar, dichotomously branched and probably radicelled
at the base. Zooecia dimorphic, the marginal zooecia of the acute edges of the
branches being larger, and having larger opesiae than the normal zooecia. Crypto-
cyst well developed, finely granulated; opesiae small with straight proximal rim
and occasionally developed lip. Distal interzooecial asymmetrical vibracula above
the opesiae with long elliptical opening and a small thornlike process going out from
the left or right inner margin. Ovicells inconspicuous exteriorly, deeply immersed,
endozooecial.
Type species: Biflustra radula Marsson 1887, Lower Maastrichtian Riigen
(Germany) .
REMARKS. This new genus comprises three characteristic species in the Maastrich-
tian, which cannot be attributed to any other genus hitherto known. Biflustra
radula Marsson was assigned incorrectly to Amphiblestrum by Voigt (1930). It
differs from Amphiblestrum not only in its bilaminar and apparently radicelled
zoarium with dimorphic zooecia and semicircular opesia in which a proximal lip is
originally developed, but also by its avicularian-like vibraculum. This has no
pivot for the articulation of the mandibula as in true avicularia, but asymmetrical
curved thorn-like processes on the inner left or right edge of the vibracula (Text-
fig. 2) and an ellipsoidal cavity in the proximal part adapted for the motion of the
seta of the vibraculum. These structures are very well shown in the type-species,
R. radula (Marsson) (pi. 3, figs. 11-12), while they are indicated in R. minor only
in some better preserved specimens.
Owing to the loss of the Marsson Collection during the last war, a neotype for
R. radula Marsson should be erected. The specimen of Biflustra radula figured
here could represent a good neotype, but it was collected from a chalk block in
glacial drift and therefore a topotype from the Lower Maastrichtian of Riigen
would be preferred.
It is difficult to decide if this genus should be regarded as belonging to the Mala-
22
UPPER CRETACEOUS POLYZOA FROM COTENTIN
costega or Coilostega. The cryptocyst is strongly calcified and the opesia is so
small that the Malacostega level has probably been exceeded. Opesiules are absent
as in many Coilostega. Nevertheless Marsson described it as Biflustra, and perhaps
he was right to assign it to the Membranimorphs, although there are many inter-
mediate forms between these two groups. This genus seems to be rather isolated
and it is difficult to attribute it to one particular family of the Malacostega.
FIG. 2. Radulopora radula (Marsson). Some zooecia and vibracularia. x6o.
12. Radulopora minor sp. n.
(PI. 3, figs. 6-io)
Holotype Upper Maastrichtian, St. Pietersberg near Maastricht (Netherlands).
0.49844. Voigt Collection.
DERIVATIO NOMINIS. The name refers to the smaller zooecial dimensions in
contrast to the type-species R. radula (Marsson) (pi. 3, figs. 11-12).
DIAGNOSIS. Radulopora with slender dichotomous branches 0-7-1 -2 mm. wide,
consisting of about three to seven alternating transverse rows of zooecia about
0-5-0 -6 mm. long. The edges of the zoarium are formed by the larger marginal
zooecia: other zooecia not clearly distinct at their margins, elongate and pyriform
with distal raised margin and a well developed cryptocyst deeply immersed proxi-
mally. Opesiae rounded quadrangular or high-semicircular rounded distally and
truncated proximally, showing in some specimens a well developed proximal lip.
Distal vibracula small, the peak turned obliquely downwards but symmetrically
oriented outwards from the median line toward the edges of the branches. Ovicells
form inconspicuous slight swellings above the opesiae (pi. 3, fig. 9).
DESCRIPTION. The bilaminar fronds have narrow cylindrical or prismatic basal
rods which are pointed toward their proximal ends and suggest an articulated basal
attachment of the zoarium. The zoarial and opesial dimensions are smaller in these
proximal parts of the zoarium than in the more distal branches. The shape of the
opesia varies from nearly semi-circular to high-oval, trapezoidal or oval: if it is
oval it is narrowed proximally but it is never circular. The straight proximal
edge of the opesia is deeply immersed in old zooecia and may disappear. The distinct
UPPER CRETACEOUS POLYZOA FROM COTENTIN 23
proximal lip observed regularly in some specimens has its origin from this straight
proximal edge. The opesia occupies about one quarter to one fifth of the length
of the zooecium. The cryptocyst is highly calcified and often appears in the proximal
part. The marginal zooecia appears to be larger than they actually are because
they are not narrowed in their proximal part, and the opesiae are always distinctly
larger than in the other zooecia, although there may be transitions between the size
of the opesiae of the normal and those of the marginal zooecia. The ovicelled
zooecia never have vibracula. The ovicells are very flat swellings above the opesiae.
If their roof is broken away a large deep hollow is revealed showing that the ovicell
is deeply immersed although it must be regarded as endozooecial.
A minute fragment of eight zooecia belongs to this species which corresponds very
well with the abundant material of the author's collection from the Cotentin localities
and from Maastricht.
MEASUREMENTS.
Lz (= Distance between the opesia) : 0-50-0-67 mm.
ho = 0-10-0-12 mm. ""I
> normal zooecia.
lo = 0-075-0-12 mm. J
ho = 0-14-0-17 mm. ~\ . ,
, > marginal zooecia.
lo = 0-15-0-16 mm. /
STRATIGRAPHICAL RANGE. Upper Maastrichtian.
SPECIMENS. 0.49579. A small worn fragment. From matrix of D.iSi.
Upper Maastrichtian, Cotentin, Manche, France.
0.49580-81. Two fragments. Upper Maastrichtian, Chef du Pont, Cotentin,
Manche, France. Voigt Collection.
0.49582. Fragment with ovicelled zooecia. Upper Maastrichtian, Cotentin,
Manche, France. Voigt Collection.
Genus HAGENOWINELLA Canu 1900
13. Hagenowinella cf. incrassata (d'Orbigny)
(PI. 2, figs. 1-3)
1853 Flustrellaria incrassata d'Orbigny: 527, pi. 726, figs. 5-8.
A large unilaminar fragment consisting of more than twenty zooecia may belong
to this species, although the opesiae are oval and not so broad and truncated as in
d'Orbigny 's type-specimen from Sainte-Colombe (Manche). A fragment of this
form collected by the author near Chef du Pont (Manche), shows, however, that the
shape of the opesiae can vary considerably and that in certain parts of the zoarium
the opesiae are fairly oval as in 0.49583, in which two zooecia have the cryptocyst
broken giving the false appearance of avicularia. The hyperstomial ovicells shown
on the specimen from Chef du Pont are broad and low. The horse shoe-like lamella
in the interior of the zooecia seems to be hidden by a fine calcitic crust, but it is
indicated in pi. 2, figs. 2-3, by two small knob-like processes at the inner margin of
the opesia.
24 UPPER CRETACEOUS POLYZOA FROM COTENTIN
SPECIMENS. 0.49583. A small fragment of zoaiium, embedded in matrix of
D.iSi. Upper Maastrichtian, Cotentin, Manche, France.
0.49845. Fragment of zoarium. Upper Maastrichtian, Fresville, Cotentin,
Manche, France. Voigt Collection. Fragment with ovicelled zooecia. Upper
Maastrichtian, Chef du Pont, Cotentin, Manche, France. Voigt Collection, Ham-
burg, Nr. 3909.
STRATIGRAPHICAL RANGE. Upper Maastrichtian.
Genus BACTRELLARIA Marsson 1887
14. Bactrellaria rugica Marsson
(PI. 2, figS. 9-14)
1887 Bactrellaria rugica Marsson 59, pi. 5, fig. 18.
1930 Bactrellaria rugica Marsson; Voigt: 444, pi. 12, figs. 25-26.
The species is represented by one very poor fragment showing four zooecia of the
frontal face with worn avicularia forming cavities between the opesiae. Although
the specimen is very small and worn, the species is so characteristic that there is
no doubt about the identification. It has been found by the author at Maastricht
and Port Filiolet (Manche) where it is rare. The frontal avicularium below the
opesia is, if well preserved, rather prominent, elongate beak-like and is proximally
oriented with a raised spatulate peak. Marsson's figure showing only a round
pore is inadequate, but he makes mention of the tube-like avicularia which occasion-
ally occur on the lateral edges of the zoaria. The frontal avicularium below the
opesia, is, if present and well preserved, rather prominent, tube-like, elongate and
proximally oriented. Its beak is dilated and spatulate but is mostly broken away
or damaged. It is rather well preserved on a specimen from an Upper Maastrichtian
chalk-bearing erratic flint-boulder found in the gravel-pit of Wulmstorf near Harburg
(pi. 2, fig. 13). If the dilated spatulate avicularian beak is destroyed there remains
an oval ring or a scar like the ones in many specimens of the Cotentin (pi. 2, figs. 9
and 12). The small pit on Marsson's figure, however, is not the cicatrix of a destroyed
avicularium, but corresponds to the spot where the avicularium is usually developed.
Three pairs of oral spines are present in well preserved specimens. The ovicells are
hyperstomial. Many fragments of the band-like zoaria from the Cotentin localities
have four to six rows of zooecia instead of three; their diameter varies between
0-7-2-0 mm. The length of the zooecia is between 0-6-0-8 mm., that of the opesia
c. 0-35 mm. In addition to the band-like specimens there occur, at Maastricht,
prismatic vincularian rods with five or more rows of zooecia showing absolutely the
same features and size of zooecia, avicularia and ovicells. They are very like
Pithodella and it is possible that they represent another mode of growth of the same
species. Eschara gaimardi von Hagenow (1851 : 82, pi. 12, fig. 10) is thought to
be an Eschara-like stem development of this genus.
STRATIGRAPHICAL RANGE. Maastrichtian.
SPECIMENS. 0.49584. A small worn fragment. From matrix of D.iSi.
Upper Maastrichtian, Cotentin, Manche, France.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 25
0.49585. A well preserved fragment. Upper Maastrichtian, Port Filiolet,
Cotentin, Manche, France. E. Voigt Collection. A worn fragment. Upper Maas-
trichtian, Port Filiolet, Cotentin, Manche, France. Voigt Collection, Hamburg
Nr. 3914.
A well preserved fragment. Upper Maastrichtian Chalk from Flint-boulder in
Pleistocene Drift. Wulmstorf near Harburg-Hamburg, Germany. Voigt Collection
Nr. 4146.
Genus STAMENOCELLA Canu & Bassler 1917
15. Stamenocella marginata (d'Orbigny)
(PI. 3, figs. 1-5)
1852 Flusirella marginata d'Orbigny: 295, pi. 700, figs. 7-9.
1852 Flustrella convexa d'Orbigny: 290, pi. 699, figs. 1-3.
1852 Biflustra tesselata d'Orbigny: 271, pi. 694, figs. 7-9.
HOLOTYPE. A small fragment of Flustrella marginata d'Orbigny. Upper Maas-
trichtian, Sainte Colombe, Cotentin, Manche, France. In d'Orbigny Collection,
Paris, Musee d'Histoire Naturelle, Nr. 8134.
A worn fragment of this species embedded in the matrix of Onychocella bathonica
Gregory (D . 181) agrees in all essential characteristics with the two specimens which
were described by d'Orbigny as Flustrella marginata from Sainte-Colombe and
Biflustra tesselata from Nehou, and which are figured here for comparison. The
latter is a worn specimen of Stamenocella marginata showing traces of ovicells and
represents the same stage of preservation as figs. 3 and 8 of pi. 31 of Canu & Bassler
(1920) where the ovicells and the avicularia of Stamenocella mediaviculifera and
Stamenocella inferaviculifera cause two shallow cavities between the opesiae.
PI. 3, fig. 5, shows a rather well preserved specimen which seems to be intermediate
between Flustrella marginata d'Orbigny and Flustrella convexa d'Orbigny. The
basal region, which tapers proximally and shows a radicelled base, has zooecia cor-
responding to those of Flustrella convexa. They are closed by a calcareous lamella
and pierced by elliptical or round openings as can also be observed in other species
of Stamenocella.
The variation in the size of the opesiae is enormous in this species as shown in
pi. 3, fig. 5, and the following measurements:
LZ = 0-50-0-63 mm.
lz = 0-17-0-25 mm.
L0 = 0-17-0-33 mm.
10 = 0-07-0-15 mm.
STRATIGRAPHICAL RANGE. Maastrichtian.
SPECIMENS. 0.47322. A fragment embedded in the matrix of Onychocella
bathonica Gregory (D.iSi). Upper Maastrichtian, Cotentin, Manche, France.
D . 49586. A fragment from the matrix of D . 181. Upper Maastrichtian, Cotentin,
Manche, France.
26 UPPER CRETACEOUS POLYZOA FROM COTENTIN
0.49587. A small worn fragment. Locality and horizon as above. A well
preserved fragment showing the stage of Flustrella convexa d'Orbigny at the base of
the zoarium. Upper Maastrichtian, Chef du Pont, Cotentin, Manche, France,
Voigt Collection, Hamburg, Nr. 4128.
Type specimen of Biflustra tesselata d'Orbigny. Upper Maastrichtian, Nehou,
Cotentin, Manche, France. d'Orbigny Collection, Paris, Musee d'Histoire Naturelle.
Nr. 8053.
Genus THYRACELLA Voigt 1930
16. Thyracella cf . meudonensis (d'Orbigny)
(PI. 2, figs. 4-6)
cf. 1851 Biflustra meudonensis d'Orbigny: 263, pi. 692, figs. 4-6.
1951 Thyracella cf. meudonensis (d'Orbigny) Voigt: 59, pi. 9, figs. 4-5.
Three fragments of this bilaminar species are conspecific with a " Biflustra "
which is very abundant in the Maastrichtian of Port Filiolet (Manche). It has a
very prominent large avicularium as is typical of Thyracella (pi. 2, fig. 4). I think
it could be identified with Biflustra meudonensis d'Orbigny recorded by him from
Ne"hou and Meudon. In the catalogue of the d'Orbigny collection only one fragment
from the Chalk of Meudon is registered under Nr. 8090 although I possess more
than fifty specimens from this locality which would fit very well into that species.
It is worn and does not correspond with the figure; it does not look like a chalk
fossil from Meudon, but seems to have come from a more littoral facies resembling
the Maastrichtian of the Cotentin. The worn specimens, described and figured by
the present author from the Maastrichtian of Kunrade (S-Limburg) and Ilten
(northern Germany) give a different impression from that of the well preserved
material from the Cotentin localities in the author's collection. But there are
all intermediate stages. For comparison a photograph of a characteristic specimen
with a large avicularium, from Port Filiolet (Manche), is given (pi. 2, fig. 4). The
length of the avicularium is c. 0-8 mm., and that of the zooecia c. 0-5-0-6 mm.
The well preserved zooecia always have a sharp and distinct margin in the distal
part as shown in the figure of d'Orbigny.
STRATIGRAPHICAL RANGE. Maastrichtian and (fide d'Orbigny) Upper Campanian.
SPECIMENS. D .49588. A damaged fragment from the matrix of D . 181. Upper
Maastrichtian, Cotentin, Manche, France.
0.49846. A fragment embedded in matrix of D.iSi. Upper Maastrichtian,
Cotentin, Manche, France.
0.49589-90. Two fragments from the matrix of D.iSi. Upper Maastrichtian,
Cotentin, Manche, France.
A well preserved branched fragment with an avicularium. Upper Maastrichtian,
Port Filiolet, Cotentin, Manche, France. Voigt Collection, Hamburg. Nr. 3919.
A worn fragment labelled Meudon but likely. Upper Maastrichtian, locality
uncertain (? Nehou, Cotentin, Manche). d'Orbigny Collection, Paris, Musee d'Histoire
Naturelle Nr. 8040.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 27
Genus VINCULARIA (auct.)
The genus Vincularia Defrance is here understood in the sense of the older authors,
although this is not correct according to the rules of nomenclature. Vincularia is
now restricted to those forms which Canu 1907 named Heterocella. But Vincularia
fragilis Defrance 1829 from the Eocene is the type species of the genus of Vincularia
(cf. Bassler 1953 : 157) and the name Vincularia must be reserved for this group.
Heterocella, therefore, as a synonym of Vincularia, must be dropped.
The consequence of this is that there is no name available for the many Cretaceous
species of " Vincularia " of d'Orbigny, Marsson, Brydone and other authors. Canu
(1900) has united most species of Vincularia under Smittipora Jullien (1881), which
Bassler (1953) considered con-generic with Diplopholeos , Rectonychocella and Velu-
mella Canu & Bassler. But the definition of Smittipora does not fit most " Vin-
cularias ". Admitted that Vincularia has been established primarily as a zoarial
growth-form for rod-like stems, there still exists a natural group of forms for which
this name has been used and for which another name does not yet exist. On the
other hand it seems to be necessary to distinguish the forms which are radicelled
or articulated at their base from those which are attached by an encrusting base.
This is not the place to give a new classification of Vincularia, and it is necessary
to retain the old name provisionally.
17. Vincularia canalifera von Hagenow
(PI. 5, figs. 7-10 and PI. 8, figs. 9-12)
1851 Vincularia canalifera von Hagenow: 61, pi. 6, fig. 14.
?i85i Vincularia flexuosa d'Orbigny: 76, pi. 656, figs. 16-18.
1930 Vincularia canalifera von Hagenow; Voigt, 467, pi. 17, fig. 18.
More than thirty fragments of this species were found in the matrix of D.iSi.
They are conspecific with the most common Vincularia-species of Maastricht and
Kunrade (Netherlands) which was described by von Hagenow as Vincularia canali-
fera. Von Hagenow's figure is not quite typical because the length of the opesia
is relatively large, about one third or one quarter of the length of the zooecium.
Among some hundred fragments there are only two which correspond to von Hage-
now's figure, but this species can show great variation in size and shape of the
zooecia and opesiae. The opesia varies between an oval (pi. 5, figs. 7, 9, 10) or
more semicircular (pi. 5, fig. 8, pi. 8, figs. 9-11) opening which is truncated more or
less proximally. In well preserved specimens it has a slight margin, but never a
proximal lip. In the proximal basal region the zooecia are 0-25 mm. long, and when
fully developed 0-5 mm. ; the length of the opesiae is 0-10-0-17 mm. The avicularia,
which have not been figured until now, reach the length of the zooecia or exceed it
(pi. 5, fig. 8, pi. 8, figs. 10, n). They are rare, straight, and appear at the beginning
of a new row of zooecia, or are normally enclosed in these. They are broader than the
autozooecia and have a flatly rounded prominent distal rim and a small elliptical
28 UPPER CRETACEOUS POLYZOA FROM COTENTIN
opesia which is longitudinally oriented. Common to the zooecia and the avicularia
is a furrow-like deepening in the median axis of the cryptocyst from which the
specific name is derived. Ovicells have never been observed.
The rods have a diameter of 0-5-0 -7 mm. and consist of eight to fourteen rows
of zooecia. They taper proximally and were articulated or radicelled at their base.
This is shown by some of the earliest zooecia which seem to be modified to lodge
chitinous rootlets (pi. 5, fig. 7). Their opesiae are smaller and their upper half is
covered by a bent calcitic lamella as is seen also in other articulated or radicelled
forms.
This species is also common in the Maastrichtian of the Cotentin near Port Filiolet,
Fresville and Chef du Pont from which region d'Orbigny described some very similar
vincularian species.
I take it that Vincularia flexuosa d'Orbigny is conspecific with this species, which
is cited by d'Orbigny from Nehou in the Cotentin and from the Santonian of Ven-
dome. I have studied the type (Nr. 7752 in the d'Orbigny collection) and I cannot
find any significant differences. It is figured on pi. 8, fig. 14 for comparison with
specimens from Maastricht (pi. 5, figs. 7 and 8 and pi. 8, figs, n and 12). The reason
why I hesitate to place this species under d'Orbigny's name is, that according to the
label the type specimen is from the Santonian of Vendome (as stated also in his
catalogue) although I suspect that it comes from Nehou in the Cotentin. Because
it is not clear if the holotype comes from the Cotentin, I prefer the name Vincularia
canalifem which is given by von Hagenow in the same year as d'Orbigny's name.
Another form from the Cotentin which is very similar to Vincularia canalifera
is V. concinna (d'Orbigny 1851 : 79, pi. 657, figs. 10-12). It may be another syno-
nym. D'Orbigny's holotype is figured on pi. 8, fig. 13; this is the only specimen
of this species in the d'Orbigny Collection of Nr. 7756). This specimen, recorded by
Canu (1900 : 420) as "insuffisant ", has opesiae also which are about 0.11-0.17 mm.
long, but the cryptocyst seems to be less deepened than in Vincularia canalifera von
Hagenow. Unfortunately it shows no avicularia, knowledge of which is very import-
ant for the discrimination of many species of Vincularia which are very similar
and cannot be identified from the work of d'Orbigny. D'Orbigny never figured
or mentioned in his species the avicularia, which are rather rare and similar to the
autozooecia.
STRATIGRAPHICAL RANGE. Maastrichtian.
SPECIMENS. 0.49732. Branched fragment, Upper Maastrichtian, Geulem near
Berg, Geul- Valley near Maastricht, Netherlands. Voigt Collection.
D. 49733, 0.49735-36, 0.49737-66. Fragments from the matrix of D.iSi,
Upper Maastrichtian, Cotentin, Manche.
0.49734. Fragment, Upper Maastrichtian Chef du Pont, Cotentin, Manche.
Voigt Collection.
0.49840-41. Two fragments, Upper Maastrichtian, St. Pietersberg near Maas-
tricht, Netherlands. Voigt Collection.
0.49842. Fragment, Upper Maastrichtian, St. Pietersberg near Maastricht,
Netherlands. Voigt Collection.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 29
Genus QUADRICELLARIA d'Orbigny 1850
18. Quadricellaria elegans d'Orbigny
(PI. 6, figs. 8-10)
1851 Quadricellaria elegans d'Orbigny: 33, pi. 652, figs. 1-5.
1900 Quadricellaria elegans d'Orbigny; Canu: 413.
1928 Quadricellaria excavata d'Orbigny; Voigt: 112, text-figs. 1-5.
1930 Quadricellaria excavata d'Orbigny ; Voigt : 489, pi. 25, figs. 12-14.
Two small fragments of a Quadricellaria may be determined as Q. elegans d'Orbigny.
The size of the opesia compared with the length of the zooecia varies considerably,
and Canu may be right in uniting d'Orbigny's three species, Q. elegans, Q. excavata
and Q. pulchella, under the name Q. excavata, as I did in 1928. My specimens agree
largely with the figures of Q. elegans d'Orbigny which was first recorded from Nehou
(Manche) .
This species is represented in my material from Chef du Pont and Port Filiolet.
One specimen from Chef du Pont has, on the narrow sides of the segments, two
enormous avicularia which are I mm. long and have an opesia with a length of
0-3 mm. (pi. 6, fig. 10).
SPECIMENS. 0.49591-92. Two worn fragments. From matrix of D.iSi.
Upper Maastrichtian, Cotentin, Manche, France.
A fragment with a large avicularium. Upper Maastrichtian, Chef du Pont,
Cotentin, Manche, France. Voigt Collection, Hamburg, Nr. 4133.
Genus COSCINOPLEURA Marsson 1887
19. Coscinopleura sp.
Two minute indeterminable fragments of a Coscinopleura, showing only a few
zooecia are present. They possibly belong to a species of Coscinopleura with small
zooecia, like Coscinopleura lamourouxi von Hagenow, or to a similar form. Common
in the Cotentin Maastrichtian near Port Filiolet and Chef du Pont (Manche).
SPECIMENS. 0.49593-94. Two small worn fragments. From the matrix of
D.iSi. Upper Maastrichtian, Cotentin, Manche, France.
Genus SEMIESCHARINELLA d'Orbigny 1852!
20. Semiescharinella complanata d'Orbigny
(PI. 4, figs. 11-15)
1840 Cellepora ricata von Hagenow: 616.
1852 Semiescharinella complanata d'Orbigny: 427, pi. 714, figs. 1-4.
1 D'Orbigny published his genera Semiescharinella and Reptescharinella on p. 427 and p. 428 of his
work. According to Sherborn (Geol. Mag. 1889 : 223-225) pages 187-472 appeared in 1852. I follow
here the dates given by Sherborn, although Lang (1917 : 172) states 1853 for Reptescharinella and
d'Orbigny himself writes 1851 for both genera.
3o UPPER CRETACEOUS POLYZOA FROM COTENTIN
1852 Escharinella simplex d'Orbigny: 205, pi. 683, figs. 14-16.
1900 Rhagasostoma simplex (d'Orbigny); Canu: 433.
1930 Micropora subgranulata (von Hagenow) Voigt (pars): 476, pi. 24, fig. 18 (non 19).
1959 Semiescharinella complanata (d'Orbigny) Voigt: 54, pi. 6, fig. i.
1962 Semiescharinella complanata (d'Orbigny) Berthelsen: 134, pi. 14, figs. 2-3.
Two very small unilaminar fragments agree very well with some specimens from
Chef du Pont and with d'Orbigny's type-specimen from Sainte Colombe (Manche).
The size of the zooecia is a little less in the type-specimen (about 0-56-0-60 mm.)
compared with 0-70 mm. in 0.49595-96; but the abundant material in the author's
collection shows that the size of the zooecia and opesiae is highly variable in this
species. The " TYPE " is a fragment, with c. 18 zooecia, in which the relative
length and width of the zooecia differ from d'Orbigny's figure as noted by Canu
(1900 : 421), although in the catalogue of the d'Orbigny Collection only one speci-
men from Sainte Colombe (Manche) is registered. The bilaminar Escharinella
simplex d'Orbigny from Nehou (Manche) belongs to the same species. The distal
pore which was interpreted by Canu as the trace of the ovicell is a true avicularium.
Nevertheless, there are some real ovicells which are developed in place of the avi-
cularia (see pi. 4, fig. 15).
Comparison of the photographs (pi. 4, figs. 13-15) shows that there is no variation
in the shape or size of the zooecia, nor of the opesiae, nor amongst the distal avicu-
laria. I have figured the only poor fragment from the d'Orbigny collection which
must be regarded as the type specimen (pi. 4, fig. 15). This species is congeneric
with Cellepora (Discopora) subgranulata von Hagenow (1851 : 91, pi. n, fig. 15)
which was chosen by Lang (1917 : 172) as the type species for the genus Repteschari-
nella d'Orbigny 1852. D'Orbigny had united under this name eight Cretaceous,
one Tertiary and two Recent species, of quite distinct systematic differences, said
to be characterized by an " ouverture mediocre ", a " pore special " and encrusting
zoaria. The genus Semiescharinella was not discussed by Lang, being represented
only by Semiescharinella complanata d'Orbigny. There is no doubt what d'Orbigny
meant by this name, and I prefer it, proposing to drop Reptescharinella as a synonym
of Semiescharinella, the mode of growth being no generic criterion. This form was
first described by von Hagenow 1840 under the name Cellepora ricata without any
figure (Voigt 1959 : 54).
STRATIGRAPHICAL RANGE. Maastrichtian-Danian.
SPECIMENS. 0.49595-96. Two small worn fragments. From the matrix of
D.iSi. Upper Maastrichtian, Cotentin, Manche, France.
0.49847. A small fragment. Upper Maastrichtian, Chef du Pont, Cotentin,
Manche, France. Voigt Collection.
A small fragment showing an ovicelled zooecium. Upper Maastrichtian, Chef
du Pont, Cotentin, Manche, France. Voigt Collection Hamburg Nr. 39i8b.
Bilaminar fragment, Holotype of Escharinella simplex d'Orbigny. Upper Maas-
trichtian, Nehou, Cotentin, Manche, France. d'Orbigny Collection, Paris, Musee
d'Histoire Naturelle Nr. 7942.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 31
Genus ROSSELIANA Jullien 1888
21. Rosseliana thomasi sp. n.
(PL i fig. 6, and PL 4, figs. 4-6)
TYPE SPECIMEN. 0.49597. Zoarium encrusting a branched fragment of Radu-
lopora minor n. g. n. sp. Upper Maastrichtian, Chef du Pont, Cotentin, Manche,
France. Collection E. Voigt.
DERIVATIO NOMINIS. In honour of Dr. Dighton Thomas, London, who first
detected this species in the matrix of Castanopora jurassica (Gregory) .
DIAGNOSIS. A Rosseliana with encrusting zoarium. Zooecia very small, oval,
only 0-33 mm. long, with smooth cryptocyst and sharply raised rim in the distal
region of the zooecia. Opesiae semicircular, occupying nearly a third of the length
of the zooecia, sometimes with very inconspicuous lateral processes, but never
trifoliate in shape; its lower rim is straight without opesiules. Ovicells globular
swellings above the opesia and occupying the proximal part of the distal zooecium.
DESCRIPTION. In addition to the British Museum specimen, which encrusts a
branch of Multicrescis laxata d'Orbigny, there are two others from Chef du Pont
(Manche). They show, although they are more fragile, the same essential specific
characters, having no avicularia and no trifoliate opesiae. In some zooecia, very
minute lateral processes might indicate a Floridina-like opesia, but this is so negli-
gible that it can be ignored. They are therefore classified as Rosseliana and not
Floridina or Floridinella. Nevertheless, it is clear that the difference between
Rosseliana and Floridinella, based only on the existence of broad opesiular indenta-
tions in the latter, is slight. The specimens are very similar to that figured by
Bassler (1953, fig. 130.4), but he only records Rosseliana from the Oligocene to
Recent.
There are very few species with which this form could be compared. The shape
and size of zooecia are very similar in Floridina (or better Floridinella) scutata Levinsen
(1925 : 345, pi. 4, fig. 39) from the Danish Maastrichtian Chalk and Danian, and
Semieschara complanata d'Orbigny (1852 : 369, pi. 708, figs. 5-8). Apart from the
free unilaminar fronds of the latter, it is very difficult to find any constant difference
between these two forms, which are distinguished from our new Rosseliana thomasi
only by their opesiae, always markedly trifoliate as in Floridina or Floridinella.
PL 4, fig. 7, shows a specimen of Floridinella scutata Levinsen from a Maastrichtian
flint drift boulder from northern Germany. The difference of this species from
Rosseliana thomasi n. sp. is clearly seen in the opesia which has significant lateral
indentations.
STRATIGRAPHICAL RANGE. Upper Maastrichtian.
SPECIMENS. 0.47323. Encrusting zoarium with some ovicelled zooecia on
Multicrescis laxata d'Orbigny (= 0.49575). In the matrix of Castanopora
jurassica (Gregory) (D . 180) Upper Maastrichtian, Cotentin, Manche, France.
Zoarium encrusting an echinoid fragment. Upper Maastrichtian, Port Filiolet,
Cotentin, Manche, France. Voigt Collection, Hamburg, Nr. 3932.
32 UPPER CRETACEOUS POLYZOA FROM COTENTIN
Genus MICROPORA Gray 1848 (non Eichwald 1855)
22. Micropora transversa (d'Orbigny)
(PI. 7, figs. 9-12)
1851 Vincularia transversa d'Orbigny: 78, pi. 657, figs. 7-9.
1887 Vincularia rugica Marsson: 65, pi. 6, fig. 8.
1930 Micropora rugica (Marsson) Voigt: 472, pi. 21, fig. 20.
1951 Micropora rugica (Marsson) Voigt: 63, pi. 9, fig. 10 and pi. 10, fig. 4.
HOLOTYPE. Two minute fragments evidently 2 pieces of one specimen. Upper
Maastrichtian, Nehou, Cotentin, Manche, France. d'Orbigny Collection, Paris,
Musee d'Histoire Naturelle Nr. 7755.
There is only a very small fragment corresponding to the length of a single zoo-
ecium of this species in the matrix of D . 181. The two symmetrical opesiules below
the opesia show clearly that it must belong to Vincularia transversa d'Orbigny, the
type of which, from Nehou, (pi. 7, figs. 10-11) was studied by the author. There is
only one poorly preserved fragment in d'Orbigny's collection which is recorded in his
catalogue. It is now broken into two pieces. It does not correspond to d'Orbigny's
figure and description, because the paired opesiules, which are clearly visible,
are neither mentioned in his text nor figured in his drawing ; they seem to be indicated
by the deep furrows accompanying the thick margin of the zooecia, but they do not
correspond to what d'Orbigny called " une depression lanceolee " which is situated
below the aperture. This depression must correspond to the lanceolate cryptocyst
of d'Orbigny's figure. I found some well preserved fragments at Port Filiolet
and Chef du Pont (Cotentin), see pi. 7, fig. 12.
This form is conspecific with Marsson's Vincularia rugica as is proved by many
specimens from the Maastrichtian Chalk of Rugen and other localities in northern
Germany. Marsson who published the first good description and figure of this
species, was therefore unable to recognize that his species was the same as d'Orbigny's,
and it is understandable that, since Marsson's description, this form has been re-
corded only under the name rugica. Another similar species is Vincularia undata
d'Orbigny from the Santonian of Vendome, which also has true opesiules below the
opesia, not indicated in d'Orbigny's figure (d'Orbigny 1851 : 75, pi. 656, figs. 10-12)
but which can be seen on the holotype.
The appearance of Micropora transversa can vary considerably as shown in
Voigt's figures (1951). The diameter of the rods is from 0-5-0 -7 mm., and the
number of zooecial rows varies between five and ten. The length of the normal
zooecia is o -49-0 -54 mm. The basal attenuated part of some rods shows clearly
that this species was articulated or radicelled (cellarif orm) . This means that this
form is not congeneric with all true Micropora, and it could be suggested that it
belongs to a new genus. But I hesitate to erect a new genus for it because it is
not impossible that this form is conspecific with Dimorphostylus tetrastichtis Voigt
1928. The genus Dimorphostylus was established for articulated rods, in which the
zooecia are only developed on one side. But the size and other features of the
UPPER CRETACEOUS POLYZOA FROM COTENTIN 33
zooecia in Micropora rugica and in Dimorphostylus tetrastichus , which are often
associated together in the same localities, are identical, and there is the suspicion
that Dimorphostylus tetrastichus might represent specimens with the zooecia de-
veloped only on one side, and that the differentiation in a frontal and reversed side
may be pathological. The two forms have not yet been found united in one speci-
men, but if they were, the generic name Dimorphostylus must be applied to Micropora
transversa d'Orbigny. Therefore it is provisionally here referred to Micropora,
which also indicates that it may belong to the Microporidae.
STRATIGRAPHICAL RANGE. Maastrichtian ; Lower Maastrichtian of Riigen and
Denmark and Upper Maastrichtian of the Cotentin (Manche), Maastricht and Ilten
near Hanover.
SPECIMENS. 0.49768. A minute fragment. From matrix of D.iSi. Upper
Maastrichtian, Cotentin, Manche, France.
0.49769. A well preserved fragment. Upper Maastrichtian, Chef du Pont,
Cotentin, Manche, France. Voigt Collection.
Genus PUNCTURIELLA Levinsen 1925
23. Puncturiella cf. superba Brydone
(PL 4, figs. 8-10)
1936 Puncturiella superba Brydone: 84, pi. 40, fig. 18.
The small unilaminar fragment with about a dozen poorly preserved zooecia,
shows the cryptocyst pierced by two outer and two inner rows of pores. These are
barely visible because they are partly obscured by recrystallization of the calcite,
and the distal avicularium which is directed obliquely is very obscure. However
the identity of this specimen with a few unilaminar fragments from Fresville and
Chef du Pont is evident (pi. 4, figs. 8-9), and they are probably identical with some
very similar specimens from Maastricht which are free or encrusting, but whose
dimensions are a little larger.
The average length of zooecia from the Cotentin localities is about 0-65-0-70 mm.,
that from Maastricht 0-80-1-00 mm., although some zooecia from the two localities
are the same size, and it may be that the difference is ecological. The only form
with which 0.49598 can be identified is Puncturiella superba Brydone from the
Lower Maastrichtian Chalk of Trimingham (Norfolk), in which the zooecia are about
0.65-1 mm. long.
STRATIGRAPHICAL RANGE. Maastrichtian.
SPECIMENS. 0.49598. A small worn fragment embedded in matrix of D.iSi.
Upper Maastrichtian, Cotentin, Manche, France.
0.49599. A small fragment. Upper Maastrichtian, Chef du Pont, Cotentin,
Manche, France. Voigt Collection.
GEOL. 17, I. 3
34 UPPER CRETACEOUS POLYZOA FROM COTENTIN
0.49600. A small well preserved fragment in matrix. Upper Maastrichtian
Md, St. Pietersberg near Maastricht. Voigt Collection.
Genus LUNULITES Lamarck
24. Lunulites sp.
One very small fragment which is worn and shows a few zooecia is indeterminable.
It seems to be conspecific with one of the numerous species of Lunulites represented
in the Maastrichtian of the Cotentin.
SPECIMENS. 0.49601. A very small indeterminable fragment. In matrix of
D.iSi. Upper Maastrichtian, Cotentin, Manche, France.
Genus ONYCHOCELLA Jullien 1881
25. Onychocella nysti (von Hagenow)
(PI. 6, figs. 3-4)
1851 Eschar a nysti von Hagenow: 78, pi. 9, figs. 15-17.
1930 Onychocella nysti (von Hagenow) Voigt: 459, pi. 16, figs. 14-16.
One small fragment belongs to this species which is very common at all European
Maastrichtian localities and which I found at Port Filiolet and Chef du Pont (Manche) .
It has not previously been recorded from the Cotentin Maastrichtian.
STRATIGRAPHICAL RANGE. Campanian-Maastrichtian.
SPECIMENS. 0.49602. A worn fragment. From the matrix of D.i8i. Upper
Maastrichtian. Cotentin, Manche, France.
0.49603. A well preserved fragment. Upper Maastrichtian, Chef du Pont,
Cotentin, Manche, France. Voigt Collection.
26. Onychocella cf. cepha (d'Orbigny)
(PL 6, figs. 1-2)
1851 Eschara cepha d'Orbigny: 143, pi. 670, figs. 8-10.
Two fragments of a narrow Onychocella belong to a species which is very abundant
near Chef du Pont (Manche) and which is not distinguishable from an Onychocella
from the Maastrichtian of Archiac (Gironde). Of the numerous species of Onycho-
cella described by d'Orbigny from the French Cretaceous, the only one which can be
compared with this form is Eschara cepha d'Orbigny from Roy an, although the type-
specimen has more slender zooecia and thinner rims surrounding the zooecia. Com-
parison with the type alone would suggest a different species, but when compared
with all the material from Archiac and Chef du Pont, these differences are much
diminished and identity is more justified than the foundation of a new species.
STRATIGRAPHICAL RANGE. Maastrichtian.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 35
MEASUREMENTS.
Lz = 0-48-0-55 mm.
lz = 0-30-0-37 mm.
h0 = 0-10-0-17 mm.
10 = 0-10-0-13 mm.
Lav = 0-70-0-75 mm.
SPECIMENS. 0.49604. A branched fragment. From the matrix of D.iSi.
Upper Maastrichtian, Cotentin, Manche, France.
D . 49605 . A small fragment . From the matrix of D . 1 8 1 . Upper Maastrichtian ,
Cotentin, Manche, France.
0.49606. A fragment. Upper Maastrichtian, Chef du Pont, Cotentin, Manche,
France. Voigt Collection.
27. Onychocella bellona (d'Orbigny)
(PI. 6, figs. 5-7)
1851 Eschara bellona d'Orbigny: 134, pi. 668, figs. 7-9.
1900 Rhagasostoma bellona (d'Orbigny) Canu (pars): 431.
Two fragments of this broad bilaminar species agree in all details with the material
collected by the author at Chef du Pont (Manche) and with as pecimen in d'Orbigny's
collection from Nehou (Manche). (d'Orbigny Collection Nr. 7812.)
The small difference between the diameter of the apertures in our figured specimen
and that of d'Orbigny is irrelevant because it is also shown in the specimens from
Chef du Pont. The opesiae of the fertile zooecia are a little longer than the others.
This species belongs to the group of Onychocella lamarcki von Hagenow, which is
closely allied to it; but the latter always shows more slender branches (1-2-2-0 mm.
diameter), and shorter and thicker zooecia, and the difference in size of the fertile
and non fertile zooecia is much more evident than in Onychocella bellona d'Orbigny.
It is the same with the fertile zooecia of Onychocella lamarcki von Hagenow from
Maastricht, but the zoaria of that species are always flat and lamellar.
MEASUREMENTS.
Lz = 0-60-0-70 mm.
lz = 0-33-0-45 mm.
h0 = 0-12-0-18 mm.
10 = 0-12-0-19 mm.
Lav = 0-75-1-00 mm.
lav = 0-22-0-27 mm.
This species has been found only in the Cotentin Maastrichtian.
STRATIGRAPHICAL RANGE. Upper Maastrichtian.
SPECIMENS. 0.49607. A worn fragment. From the matrix of D.iSi. Upper
Maastrichtian, Cotentin, Manche, France.
GEOL. 17, I. 3§
36 UPPER CRETACEOUS POLYZOA FROM COTENTIN
0.49608. A well preserved fragment. Upper Maastrichtian, Chef du Pont,
Cotentin, Manche, France. Voigt Collection.
A well preserved fragment with ovicelled zooecia. Upper Maastrichtian, Nehou,
Cotentin, France. d'Orbigny Collection, Paris, Musee d'Histoire Naturelle Nr. 7812.
Genus ONYCHOCELLARIA Voigt 1957
28. Onychocellaria caecilia (d'Orbigny)
(PL 5, %s. 1-6)
1851 Eschara caecilia (d'Orbigny): 138, pi. 669, figs. 4-7.
The zoarium was cellariiform in growth. This is proved by the tapering proximal
ends of the segments and by pits occasionally found on the cryptocyst in which
rootlets are inserted. This criterion, combined with the straight avicularia, puts it
in the genus Onychocellaria, although the endozooecial ovicell, which is characteristic
for this genus, has not yet been observed.
The zooecia, the length of which is 0-4-0-5 mm., are nearly rectangular and are
very often distinguished by a small horizontal band above the distal rim of the
opesia. The slit-like pit between the opesia and this band is very characteristic,
and is shown in d'Orbigny's figures. The opesiae, which vary greatly in size, may
be rounded or oval to high-semicircular. In some segments the whole opesia or its
upper half is closed by a calcareous lamella.
MEASUREMENTS.
Lz = 0-42-0-51 nun.
lz = 0-20-0-25 mm.
h0 = o-io-0'70 mm.
10 = o-io-o-n mm.
LAV = 0-55-0-58 mm.
1AV = 0-25-0-30 mm.
This species is represented by many small fragments. It is characteristic of the
Maastrichtian of the Cotentin, and has also been found in abundance by the present
author near Port Filiolet (Manche). The specimens agree entirely with the original
specimens of d'Orbigny from the Maastrichtian of Nehou. These are not " use "
as stated by Canu (1900 : 420).
STRATIGRAPHICAL RANGE. Upper Maastrichtian.
SPECIMENS. 0.49726. A small fragment with avicularia. From the matrix of
D.iSi. Upper Maastrichtian, Cotentin, Manche, France.
0.49727. Another fragment with avicularium. From the matrix of D.iSi.
Upper Maastrichtian, Cotentin, Manche, France.
D. 49728-29. Two worn fragments. From the matrix of D.iSi. Upper Maas-
trichtian, Cotentin, Manche, France.
0.49767. A small fragment corresponding in preservation to 0.49730. From
the matrix of D.iSi. Upper Maastrichtian, Cotentin, Manche, France.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 37
0.49849-54. 6 fragments from the matrix of D.iSi. Upper Maastrichtian,
Cotentin, Manche, France.
D . 49848. Eight very worn small fragments from the matrix of D . 181. Locality
and horizon as above.
D. 49730. A well preserved fragment with some avicularia. Upper Maastrich-
tian, Chef du Pont, Cotentin, Manche, France. Voigt Collection.
D. 49731. A well preserved fragment. Upper Maastrichtian, Chef du Pont,
Cotentin, Manche, France. Voigt Collection.
Genus PLIOPHLOEA Gabb & Horn 1862
29. Pliophloea sp.
(PI- 7, ng. 3)
Of this species there is but one encrusting young zoarium, which is composed
of the ancestrula and a dozen zooecia. The ancestrula is 0-20 mm. long and the
other zooecia have a length up to 0-50 mm. The smooth intraterminal front wall
shows about twelve to thirteen costae which are barely visible. The orifice is cribri-
line to slightly pliophloean according to Lang's definition. There is only one
avicularium — if the interpretation as avicularium is correct — which is c. 0-20 mm.
long and which is close to the ovicelled zooecium on the right of our figure. Details
of the spines are not visible. Pliophloea gluma Lang (1921 : 188, pi. 6, fig. 3) from
the Danian shows some affinities in the shape of the zooecia and the apertures, but
has clearly distinct costae, and visible intercostal fusions, and the zooecia are only
0-40 mm. long. There is no other species hitherto described with which this form
can be identified and therefore it may be supposed that it is a new one. But this
one poor specimen does not suffice to found a new species.
STRATIGRAPHICAL RANGE. This form has not previously been observed in the
Maastrichtian of the Cotentin.
SPECIMENS. 0.47324. Small encrusting zoarium with ancestrula and an ovi-
celled zooecium. In the matrix of Castanopora jurassica (Gregory) D . 180. Upper
Maastrichtian, Cotentin, Manche, France.
Genus DECURTARIA Jullien 1886
(= Prosoporella Marsson 1887)
30. Decurtaria cf. cornuta (Beissel)
(PI. 7, figs. 1-2)
1865 Semiescharipora cornuta Beissel: 58, pi. 7, figs. 77-81.
1887 Prosoporella cornuta Marsson : 100.
1922 Decurtaria cornuta (Beissel) Lang: 388, text-fig. 124, pi. 8, fig. 9.
1925 Barroisina trifossata Levinsen: 387, pi. 8, fig. 6.
1930 Decurtaria cornuta (Beissel) Voigt: 516, pi. 32, fig. 6.
38 UPPER CRETACEOUS POLYZOA FROM COTENTIN
This species is represented by a fairly large zoarium composed of about thirty
zooecia. Unfortunately it is covered by a film of minute calcite-crystals which
hides many of the characteristic minute details of the costae and the orifices. The
small number of costae (six to seven) and the very stout distal shield indicate it to
be Decurtaria cornuta, although the shape of the orifice is more like that of Decurtaria
allecta Lang (1922 : 386, pi. 8, fig. 8) from the Upper Maastrichtian of Maastricht.
D . 49609 corresponds exactly to a fragment from Chef du Pont (Manche) . In some
respects that form might be regarded as intermediate between the two species, which
are, indeed, more alike than is stated by Lang who had only one specimen of his
D. allecta from Maastricht. Decurtaria cornuta is not confined to the Lower Maas-
trichtian as might be concluded from Lang's work. I found a small but typical
fragment of this species in the Upper Maastrichtian of Biebosch (South Limburg,
Netherlands) .
It seems that the shape of the orifice is not so constant as was assumed by Lang.
According to him, in D. cornuta the orifice should be " super-cribriline " and in D.
allecta " supernormal ". But Beissel had already stated that the different types of
orifice shape may be observed in the same zoarium, and he has figured both forms
in his figs. 77 and 78. The size of the zooecia is between 0-57 and 0-65 mm. Barroi-
sina trifossata Levinsen is a synonym of this species which I concluded from the
examination of Levinsen's type specimen.
STRATIGRAPHICAL RANGE. Maastrichtian.
SPECIMENS. 0.49609. A complete zoarium with ancestrula and some ovicelled
zooecia. From the matrix of D.iSi. Upper Maastrichtian, Cotentin, Manche,
France.
An incomplete zoarium. Upper Maastrichtian, Chef du Pont, Cotentin, Manche,
France. Voigt Collection, Hamburg, Nr. 4137.
Genus FRURIONELLA Canu & Bassler 1926
30. Frurionella fertilis sp. n.
(PI. 8, figs. 5-7)
HOLOTYPE. 0.49610. Upper Maastrichtian, Chef du Pont, Cotentin, Manche,
France. Collection E. Voigt.
DERIVATIO NOMINIS. Derived from fertilis = generative, on account of the
numerous ovicells.
DIAGNOSIS. A slender species of Frurionella, with bilaminar branches consisting
of three to four alternating rows of zooecia. Zooecia not distinct, small, with
quadrangular to high triangular or semicircular opesiae whose proximal margin is
straight and generally provided with an inconspicuous proximal lip caused by the
swelling of the median avicularium below the opesia. Elliptical or linear indistinct
pores which may be avicularia are developed in the median axis of the zooecium.
Ovicells numerous, forming deep characteristic cavities above the apertures if the
covering lamina has been destroyed.
UPPER CRETACEOUS POLYZOA FROM COTENTIN 39
MEASUREMENTS.
Lz = 0-50-0-62 mm.
h0 = 0-10-0-12 mm.
10 = o -10-0 -12 mm.
REMARKS. A small fragment, showing a scarcely constricted opesia with a
minute avicularium below it, has been recognized as belonging to this new species
which is represented from Chef du Pont by two other similar specimens. This
form cannot be identified with any of the few known species of Frurionella. It is
smaller in its zoarial and zooecial dimensions than the other species of this genus.
At first it was supposed that it might be the slender distal branch of another new
species of Frurionella which is much larger and very common at Fresville (Cotentin,
Manche). This is improbable because there are no intermediate stages between
these two very different forms and no distal branches smaller than 2-1-5 mm. seem
to exist in the second species. This other species of Frurionella from Fresville
(pi. 8, fig. 8) is conspecific with Frurionella europaea Voigt (1951 : 60, pi. 9, figs. 1-3)
from the Upper Maastrichtian of Ilten (Hannover) and Kunrade (Netherlands).
It has now also been found at Maastricht (Netherlands).
STRATIGRAPHICAL RANGE. Maastrichtian.
SPECIMENS. 0.49610. Holotype see above.
D . 49611. A small worn fragment in matrix. From the matrix of D . 181. Upper
Maastrichtian, Cotentin, Manche, France.
0.49612. A fragment with broken ovicelled zooecia. Horizon and locality as
above.
Genus BEISSELINA Canu 1913
32. Beisselina striata (Goldfuss)
(PI. 8, figs. 3-4)
1826 Eschara striata Goldfuss: 25, pi. 8, fig. 16.
1960 Beisselina striata (Goldfuss) Wiesemann: 27, pi. i, figs. 1-3, pi. 2, figs. 3-4; pi. 12, figs 3-4;
text-figs. 36, 4 9-10; 5, Nrs. 12-13 (with additional synonymy).
1967 Beisselina striata (Goldfuss) Voigt: 72, pi. 25, fig. i.
One specimen belongs to this common Maastrichtian species which is, according
to Wiesemann, represented near Chef du Pont (Manche) . Nevertheless the diameter
of the stem (1-5 mm.) and the orifices (peristomicia) are a little smaller (0-08-0-12
mm.) than in most specimens from the type locality of Maastricht, and Kunrade,
although the length of the zooecia (distance between proximal and distal apertures)
is the same. Specimens with such small orifices are not lacking, and all intermediate
sizes occur at these localities, but they are rare. For comparison see pi. 8, fig. 4,
showing the surface of a worn specimen from Maastricht itself. These small forms
are more common in the " Tuffeau de St. Symphorien " in the Basin of Mons (Bel-
gium.)
4o UPPER CRETACEOUS POLYZOA FROM COTENTIN
STRATIGRAPHICAL RANGE. Maastrichtian.
SPECIMENS. 0.49721. A worn fragment. From the matrix of D.i8i. Upper
Maastrichtian, Cotentin, Manche, France.
0.49722. A worn fragment. Upper Maastrichtian, Geulem, near Berg, Geul
valley near Maastricht (Netherlands). Voigt Collection.
Genus BEISSELINOPSIS Voigt 1951
33. Beisselinopsis flabellata (d'Orbigny)
(PL 8, figs. 1-2)
1852 Escharifora flabellata d'Orbigny: 460, pi. 715, figs. 10-12.
non 1930 Beisselina flabellata (d'Orbigny) Voigt: 525, pi. 34, fig. n.
One incomplete young zoarium represents d'Orbigny's species which is common
at Sainte Colombe (Manche) and the type of which from this locality was studied
by the present author. There are some specimens from Chef du Pont (Manche)
which agree very well with Beisselinopsis flabellata d'Orbigny although they are
more elongate than the flabelliform zoarium of d'Orbigny's type specimen. The
species identified as Porina flabellata (d'Orbigny) by Marsson (1887 : 85) is Beisselin-
opsis marginata v. Hagenow 1839 (cf. Voigt 1959 : n, pi. 9, figs. 1-2). The similar
Danian and Montian form determined as Porina or Beisselina flabellata (d'Orbigny)
by Levinsen (1925, pi. 7, fig. 83) and Voigt (1930 partim : 525, pi. 34, fig. n) is
neither conspecific nor congeneric and corresponds to Eschara oblita Kade (1852 : 29,
pi. i, fig. 18) as stated by Berthelsen (1962 : 201, pi. 24, fig. 6) and Voigt (1964 : 458,
pi. 8, fig. 8 and pi. 14, figs. 1-3) and belongs to the genus Pavobeisselina Voigt 1964.
It was formerly assigned to Beisselinopsis Voigt 1951, but this attribution was
incorrect because Beisselinopsis has no ascopore which can always be observed in
Pavobeisselina. The inner structure of Beisselinopsis flabellata d'Orbigny does not
show any ascopore in the frontal wall as is always the case in Beisselina and flabelli-
form Pavobeisselina. Therefore it must be regarded as a true Beisselinopsis.
STRATIGRAPHICAL RANGE. Upper Maastrichtian.
SPECIMENS. 0.49723. A young zoarium. From the matrix of D.iSi. Upper
Maastrichtian, Cotentin, Manche, France.
An adult zoarium. Upper Maastrichtian, Chef du Pont, Cotentin, Manche,
France. Voigt Collection, Hamburg, Nr. 3908.
II. BRACHIOPODA
Genus THECIDEA Def ranee 1832
Thecidea papillata (von Schlottheim)
(PI. i, figs. 9-10)
1959 Thecidea papillata von Schlotheim; Backhaus: 21, pi. i, figs. 1-4 (see for all references).
UPPER CRETACEOUS POLYZOA FROM COTENTIN 41
TABLE I.
List of the species from the matrix of " Castanopora " jurassica (Gregory) (D . 180)
and Onychocella piriformis (Goldfuss) (= Onychocella bathonica (Gregory) (D.iSi).
I. POLYZOA
1. Berenicea sp.
2. Entalophora benedeniana (von Hagenow) 1851
3. Idmidronea macilenta (von Hagenow) 1851
4. Heterocrisina communis (d'Orbigny) 1853
5. Crisisina carinata (Roemer) 1840
6. Osculipora truncata (Goldfuss) 1826
7. Reteporidea lichenoides (Goldfuss) 1826
8. Petalopora sp.
9. Multicrescis laxata (d'Orbigny) 1854
10. Membranipora unipora (Marsson)
11. Amphiblestrella elegans von (Hagenow) 1851
12. Radulopora minor n. sp.
13. Hagenowinella cf. incrassata (d'Orbigny) 1853
14. Bactrellaria rugica Marsson 1887
15. Stamenocella marginata (d'Orbigny) 1852
1 6. Thyracella cf. meudonensis d'Orbigny 1851
17. Vincularia canalifera von Hagenow 1851
18. Quadricellaria elegans (d'Orbigny) 1951
19. Coscinopleura sp.
20. Semiescharinella complanata d'Orbigny 1852
21. Rosseliana thomasi n. sp.
22. Micropora transversa (d'Orbigny) 1851
23. Puncturiella cf. superba Brydone 1936
24. Lunulites sp.
25. Onychocella nysti (von Hagenow) 1851
26. Onychocella cf. cepha (d'Orbigny) 1851
27. Onychocella bellona (d'Orbigny) 1851
28. Onychocellaria caecilia (d'Orbigny) 1851
29. Pliophloea sp.
30. Decurtaria cf. cornuta (Beissel) 1865
31. Frurionella fertilis n. sp.
32. Beisselina striata (Goldfuss) 1926
33. Beisselinopsis flabellata (d'Orbigny) 1852
II. BRACHIOPODA
34. Thecidea papillata (von Schlottheim) 1813
* After d'Orbigny.
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42 UPPER CRETACEOUS POLYZOA FROM COTENTIN
Two small valves of this very common and characteristic brachiopod of the
" Craie a Thecidees " were found in the matrix of D.iSi. They are indistinguish-
able from those of the Cotentin Maastrichtian collected by the author. The collec-
tion of the " Geologisches Staatsinstitut " Hamburg possesses about 850 specimens
from Chef du Pont, Fresville and Port Filiolet, which were studied in the monograph
by Backhaus. The locality " Port Fiolet " (Backhaus 1959 : 27 and Hofker 1959 :
369, 380, 381) should be " Port Filiolet ".
STRATIGRAPHICAL RANGE. Maastrichtian, mainly Upper Maastrichtian.
SPECIMENS. 66.42981. Pedicle valve. From the matrix of D.iSi. Upper
Maastrichtian, Craie a Thecidees, Cotentin, Manche, France.
66.42982. Small dorsal valve. Horizon and locality as above.
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UPPER CRETACEOUS POLYZOA FROM COTENTIN 43
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44 UPPER CRETACEOUS POLYZOA FROM COTENTIN
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EXPLANATION OF PLATES
Some of the specimens of Bryozoa and Brachiopoda referred to and figured in
this publication are housed in the collections of the British Museum (Natural History)
and are prefixed by the letters D and BB respectively.
PLATE i
Petalopora sp. (p. 18)
FIG. i. Worn branch with narrow ribs between the small mesopores. From the matrix of
D.iSi, Upper Maastrichtian (Cotentin, Manche). X25. 0.49573.
FIG. 2. Branch in similar condition to fig. i with larger mesopores and peristomes. Upper
Maastrichtian, Port Filiolet (Cotentin, Manche). Presented by the author. X25. 0.49574.
FIG. 3. Well preserved branch. Upper Maastrichtian, Chef du Pont (Cotentin, Manche).
X25- Voigt Collection, Hamburg, Nr. 3969.
Crisisina carinata (Roemer) (p. 17)
FIG. 4. Worn fragment with damaged peristomes, lateral view. From the matrix of D. 181
Upper Maastrichtian (Cotentin, Manche). X25. 0.49566.
FIG. 5. Worn fragment, frontal view. Upper Maastrichtian, Port Filiolet (Cotentin,
Manche). Presented by the author. X25. 0.49567.
Multicrescis laxata d'Orbigny (p. 18) (See also pi. 4 figs. 4-6)
FIG. 6. Worn fragment 0.49575 with encrusting Rosseliana thomasi n. sp. (0.47323) in
matrix of Castanopora jurassica (Gregory) (D.iSo). Upper Maastrichtian (Cotentin, Manche).
X20.
FIG. 7. Holotype. Upper Maastrichtian, Sainte Colombe (Cotentin, Manche) in coll.
d'Orbigny Nr. 8416, Musee d'Histoire Naturelle, Paris, x 12.
FIG. 8. Holotype. X25.
Thecidea papillata von Schlottheim (p. 40)
FIG. 9. Pedicle (ventral) valve with growth-facet on the apex, with the cast of a small
lamellibranch on the right. From the matrix of D.iSi. Upper Maastrichtian (Cotentin,
Manche), attesting the origin from the " Craie a Thecidees " of the Cotentin-region. c. X 12.
BB. 42981.
FIG. 10. Small dorsal valve slightly damaged. From the matrix of D.iSi. c. X 12.
86.42982.
Bull. Br. Mus. nat. Hist. (Geol.) 17, i
PLATE i
8
GEOL. 17, I.
4§
PLATE 2
Hagenowinella cf. incrassata (d'Orbigny) (p. 23)
FIG. i. Part of the encrusting zoarium showing an avicularium in the left upper corner and
some damaged zooecia in the proximal region. From the matrix of D. 181. Upper Maastrich-
tian (Cotentin, Manche). x 20. 0.49583.
FIG. 2. Worn specimen with broken ovicells. The deeply immersed horseshoe-like processes
in the interior of the zooecia can hardly be seen. Upper Maastrichtian, Fresville (Cotentin,
Manche). Presented by the author. X2o. 0.49845.
FIG. 3. Unilaminar fragment with large opesia. Some ovicells and an avicularium in the
lower row of zooecia are visible. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). x 20
Voigt Collection, Hamburg, Nr. 3909.
Thyracella cf. tneudonensis (d'Orbigny) (p. 26)
FIG. 4. Branched narrow fragment with an avicularium, for comparison with fig. 5. Upper
Maastrichtian, Port Filiolet, (Cotentin, Manche). x 20. Voigt Collection, Hamburg, Nr. 3919-
FIG. 5. Damaged fragment of the bilaminar zoarium. From the matrix of D.iSi. Upper
Maastrichtian (Cotentin, Manche). X2O. 0.49588.
FIG. 6. Worn fragment, d'Orbigny Collection, Paris Nr. 8040, Musee d'Histoire Naturelle.
labelled Meudon but likely Upper Maastrichtian, locality uncertain (? Neliou, Cotentin,
Manche). X2o.
" Membranipora unipora " Marsson (p. 19)
FIG. 7. Fragment with hyperstomial ovicells and median pores on the gymnocyst of the
undamaged zooecia. The minute marks of spines are hidden by recrystallization. From the
rock matrix of D.iSi. Upper Maastrichtian (Cotentin, Manche). x 20. 0.49576.
FIG. 8. Holotype, of Flustrellaria simplex d'Orbigny Collection, Nr. 8130, Musee d'Histoire
Naturelle Paris, Upper Maastrichtian, N6hou (Cotentin, Manche). x 20.
Bactrcllaria rugica Marsson (p. 24)
FIG. 9. Small worn fragment showing the pits of the avicularia below the opesium. From
the matrix of D.iSi. Upper Maastrichtian (Cotentin, Manche). X2O. 0.49584.
FIG. 10. The same specimen, showing the backside, x 20.
FIG. ii. Fragment with well preserved avicularia. Upper Maastrichtian, Port Filiolet
(Cotentin, Manche). Presented by the author, x 20. 0.49585.
FIG. 12. Worn fragment in similar condition as figs. 9-10. Upper Maastrichtian, Port
Filiolet (Cotentin, Manche). x 20. Voigt Collection, Hamburg, ^.3914.
FIG. 13. Well preserved fragment with raised avicularia whose rostra are damaged. From
chalk of a flint-boulder of Upper Maastrichtian age, Wulmstorf near Harburg-Hamburg,
Northern Germany, x 20. Voigt Collection, Hamburg, Nr. 4146.
FIG. 14. The same as fig. 13, lateral view, showing the long tubular peduncles of the avicu-
laria. x 20.
Bull. Br. Mus. nat. Hist. (Geol.) 17, i
PLATE 2
PLATE 3
Stamenocella marginata d'Orbigny (p. 25)
FIG. i . Fragment of the bilaminar zoarium in the matrix of Onychocella piriformis (Goldf uss)
(D.iSi). Upper Maastrichtian (Cotentin, Manche). X 20. 0.47322.
FIG. 2. Similar fragment. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). Pre-
sented by the author. X2O. 0.49586.
FIG. 3. Holotype of Biflustra tesselata d'Orbigny. Upper Maastrichtian, Nehou (Cotentin,
Manche). d'Orbigny Collection, Nr. 8053, Paris Musee d'Histoire Naturelle. X2o.
FIG. 4. Holotype of Biflustra marginata d'Orbigny. Upper Maastrichtian, Sainte Colombe
(Cotentin, Manche). d'Orbigny Collection, Nr. 8134, Paris Musee d'Histoire Naturelle. X 20.
FIG. 5. Well preserved large fragment showing the different aspect and size of zooecia and
avicularia. The basal region represents Flustrella convexa d'Orbigny. Upper Maastrichtian,
Chef du Pont (Cotentin, Manche). X20. Voigt Collection, Hamburg, Nr. 4128.
Radtilopora minor n. sp. (p. 22)
FIG. 6. Small worn bilaminar fragment from the matrix of D.iSi. Upper Maastrichtian,
(Cotentin, Manche). X2o. 0.49579.
FIG. 7. Fragment showing the gradual tapering of the zoarium. Upper Maastrichtian,
Chef du Pont (Cotentin, Manche). Presented by the author. X2O. 0.49580.
FIG. 8. Another fragment. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). Pre-
sented by the author. X2O. 0.49581.
FIG. 9. Broad fragment, most zooecia with ovicells on the right side with anen crusting
Foraminifer. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). Presented by the
author. X2o. 0.49582.
FIG. 10. Holotype, showing the larger marginal zooecia at the edge of the zoarium. Upper
Maastrichtian, St. Pietersberg near Maastricht (Netherlands), x 20. Presented by the author.
0.49844.
Radulopora radula (Marsson) (p. 21)
FIG. ii. Fragment of the thick bilaminar zoarium for comparison with Radulopora minor
n. sp., showing the larger zooecia and avicularia and some fertile zooecia with broken ovicells.
Upper Maastrichtian Chalk in drift, Tornesch near Elmshorn (Schleswig-Holstein, Northern
Germany). X2O. Voigt Collection, Hamburg Nr. 3974.
FIG. 12. Singular vibraculum, x 120, showing the thornlike process which is curved inwards.
Horizon, locality and collection as for fig. n.
Bull. Br. Mus. nat. Hist. (Geol.) 17, i
PLATEs
11
8
PLATE 4
Amphiblestrella elegans (v. Hagenow) (p. 20)
FIG. i. Small worn fragment from the matrix of D.iSi. Upper Maastrichtian (Cotentin,
Manche). x 20. 0.49577.
FIG. 2. Holotype of Flustrella baculina d'Orbigny. Upper Maastrichtian, N6hou (Cotentin,
Manche). d'Orbigny Collection, Paris, Nr. 8127, Musee d'Histoire Naturelle. x 20.
FIG. 3. Branched fragment. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). Pre-
sented by the author. X2O. 0.49578.
Rosseliana thomasi n. sp. (p. 31) (see also pi. i, fig. 6)
FIG. 4. Holotype, showing the encrusting zoarium upon a branched fragment of Radulopora
minor n. sp. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). Presented by the
author, x 20. 1X49597.
FIG. 5. Same specimen, part of the zoarium x 40, showing the form of the opesium and some
fertile zooecia with ovicells.
FIG. 6. Zoarium encrusting an echinoid fragment. Upper Maastrichtian, Port Filiolet
(Cotentin, Manche). X2o. Voigt Collection, Hamburg, Nr. 3932.
Floridina scut at a (Levinsen) (p. 31)
FIG. 7. Part of encrusting zoarium, for comparison with Rosseliana thomasi n. sp. Upper
Maastrichtian chalk boulder in drift, Tornesch near Elmshorn (Schleswig-Holstein, Northern-
Germany). X40. Voigt Collection, Hamburg, ^.4145.
Puncturiella cf . superba Brydone (p. 33)
FIG. 8. Small unilaminar fragment in matrix of D.iSi. Upper Maastrichtian (Cotentin,
Manche). X2O. 0.49598.
FIG. 9. Unilaminar fragment. Upper Maastrichtian, Chef du Pont (Cotentin, Manche).
Presented by the author, x 20. D . 49599.
FIG. 10. Unilaminar fragment, well preserved. Upper Maastrichtian, St. Pietersberg near
Maastricht (Netherlands) . Presented by the author, x 20. D . 49600.
Setniescharinella complanata d'Orbigny (p. 29)
FIGS. II-I2. Small badly preserved unilaminar fragment from the matrix of D. 181. Upper
Maastrichtian (Cotentin, Manche). x 20. 0.49595 and 49596.
FIG. 13. Fragment with well preserved zooecia and distal avicularia. Upper Maastrichtian,
Chef du Pont (Cotentin, Manche). Presented by the author. X2o. 0.49847.
FIG. 14. Small fragment showing one zooecium with ovicell in the right lower corner. Upper
Maastrichtian, Chef du Pont (Cotentin, Manche) . x 20. Voigt Collection, Hamburg, Nr.
39i8b.
FIG. 15. Holotype of Escharinella simplex d'Orbigny. Upper Maastrichtian, N6hou (Coten-
tin, Manche). d'Orbigny Collection, Paris, Nr. 7942, Mus6e d'Histoire Naturelle. x 20.
Bull. Br. Mus. nat. Hist. (Geol.) 17, i
PLATE4
PLATE 5
Onychocellaria caecitia (d'Orbigny) (p. 36)
FIG. I. Fragment of a well preserved segment with some avicularia on the edges of the
slightly flattened rods. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). x 20.
Presented by the author. 0.49730.
FIG. 2. Same specimen seen from the narrow side with two avicularia. x 20.
FIG. 3. Another fragment with closed zooecia in the proximal region and larger opesia.
Upper Maastrichtian, Chef du Pont (Cotentin, Manche). Presented by the author, x 20.
0.49731-
FIG. 4. Small fragment with avicularium. From the matrix of D.iSi. Upper Maastrich-
tian, Chef du Pont (Cotentin, Manche). X2O. 0.49726.
FIG. 5. Another fragment with avicularium in the middle row of zooecia. From the matrix
of D.iSi. Upper Maastrichtian (Cotentin, Manche). X2o. Photographed under water.
0.49727.
FIG. 6. Broad worn fragment. From the matrix of D . 1 81 . Upper Maastrichtian (Cotentin,
Manche). X2O. Photographed under water. 0.49767.
Vincularia canalifera (von Hagenow) (p. 27) (See also pi. 8, figs. 9-12.)
FIG. 7. Proximal part of a well preserved segment, showing more oval opesia and some
zooecia with the characteristic openings for radicell filaments at the lower end. Upper Maas-
trichtian, Md., St. Pietersberg near Maastricht (Netherlands). X 20. Presented by the
author. D . 49840.
FIG. 8. Fragment with some avicularia in the distal and proximal part, showing more
semicircular opesia than fig. 7. Upper Maastrichtian, Md., St. Pietersberg near Maastricht
(Netherlands). X2O. Presented by the author. 0.49841.
FIG. 9. Worn fragment. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). x 20.
Presented by the author. 0.49734.
FIG. 10. Worn fragment. From the matrix of D.iSi. Upper Maastrichtian (Cotentin,
Manche). x 20. 0.49733.
Onychocella pirifortnis (Goldfuss) (p. 5)
FIG. ii. Fragment of the unilaminar frond with an avicularium in the lower left corner for
comparison with fig. 12. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). x 20.
Voigt Collection, Hamburg, Nr. 3573.
FIG. 12. Holotype of Onychocella bathonica Gregory, erroneously labelled as " Bathonian
Ranville (Calvados, France) ", but certainly from the Upper Maastrichtian of the Cotentin
(Manche). X2O. D.iSi.
Bull. Br. Mm. nat. Hist. (Geol.) 17, i
PLATEs
PLATE 6
Onychocella cf. cepha d'Orbigny (p. 34)
FIG. i. Branched fragment with some avicularia from the matrix of D.iSi. Upper Maas-
trichtian (Cotentin, Manche). x 20. 0.49604.
FIG. 2. Fragment for comparison with fig. i. Upper Maastrichtian, Chef du Pont (Cotentin,
Manche). X2o. Presented by the author. 0.49606.
Onychocella nysti (v. Hagenow) (p. 34)
FIG. 3. Worn fragment from the matrix of D . 181. Upper Maastrichtian (Cotentin, Manche).
X20. 0.49602.
FIG. 4. Well preserved specimen with two ovicells. Upper Maastrichtian, Chef du Pont
(Cotentin, Manche). X2O. Presented by the author. 0.49603.
Onychocella bellona (d'Orbigny) (p. 35)
FIG. 5. Well preserved fragment with several ovicells. Upper Maastrichtian, Nehou (Coten-
tin, Manche). x 20. d'Orbigny Collection, Paris, Nr. 7812.
FIG. 6. Fragment from the Upper Maastrichtian from Chef du Pont (Cotentin, Manche)
intermediate between fig. 5 and fig. 7. Presented by the author, x 20. 0.49608.
FIG. 7. Worn fragment from the matrix of D. 181. Upper Maastrichtian (Cotentin, Manche).
X40. 0.49607.
Quadricellaria elegans d'Orbigny (p. 29)
FIG. 8. Fragment of a segment. Upper Maastrichtian (Cotentin, Manche). From the
matrix of D.iSi. x 20. 0.49591.
FIG. 9. Another worn fragment. Upper Maastrichtian (Cotentin, Manche). From the
matrix of D.iSi. x 20. 0.49592.
FIG. 10. Fragment with a large avicularium. Upper Maastrichtian, Chef du Pont (Cotentin,
Manche). X4O. Voigt Collection, Hamburg, Nr. 4133.
Bull. Br. Mus. nat. Hist. (Geol.) 17, i
PLATE6
10
PLATE 7
Decurtaria cf. cornuta (Beissel) (p. 37)
FIG. i. Zoarium with ancestrula and one ovicelled zooecium. From, the matrix of D.iSi.
Upper Maastrichtian (Cotentin, Manche). x 20. 0.49609.
FIG. 2. Incomplete zoarium with some ovicelled zooecia. Upper Maastrichtian, Chef du
Pont (Cotentin, Manche). X2O. Voigt Collection, Hamburg, ^.4137.
Pliophloea sp. (p. 37)
FIG. 3. Young zoarium with ancestrula and one ovicelled zooecium. In the matrix of
Castanopora jurassica (Gregory) D.iSo. Upper Maastrichtian (Cotentin, Manche). X2o.
D.47324.
Castanopora jurassica (Gregory) (p. 7)
FIG. 4. Bilaminar fragment with two ovicelled zooecia. Upper Maastrichtian, Chef du
Pont (Cotentin, Manche). Presented by the author. X2O. 0.49724.
FIG. 5. Same specimen from the other side, showing one zooecium with preserved frontal
wall, x 20.
FIG. 6. Fragment with some zooecia. Upper Maastrichtian, Chef du Pont (Cotentin,
Manche). x 20. Voigt Collection, Hamburg, ^.3585.
FIG. 7. Part of a young zoarium with smaller zooecia. Upper Maastrichtian, Chef du Pont
(Cotentin, Manche). Presented by the author. X2O. 0.49725.
" Membranipora " constricta d'Orbigny (p. 13)
FIG. 8. Badly preserved specimen encrusting a Brachiopod. Aptian les Croutes (Yonne,
France) ? Holotype, d'Orbigny Collection, Paris, Mus6e d'Histoire Naturelle, Nr. 5691. X2O.
Micropora transversa (d'Orbigny) (p. 32)
FIG. 9. Small worn fragment showing the two opesiular pores below the opesium. From the
matrix of D.iSi. Upper Maastrichtian (Cotentin, Manche). x 20. 0.49768.
FIGS. lo-n. Two fragments of the Holotype. Upper Maastrichtian, Neliou (Cotentin,
Manche). X2O. d'Orbigny Collection, Paris, Musee d'Histoire Naturelle, Nr. 7755.
FIG. 12. Fragment, Upper Maastrichtian, Chef du Pont (Cotentin, Manche). X2O. Pre-
sented by the author. 0.49769.
Bull. Br. Mus. nat. Hist. (Geol.) 17, i
PLATE7
PLATE 8
Beisselinopsis flabellat a (d'Orbigny) (p. 40)
FIG. i. Adult zoarium. Upper Maastrichtian, Chef du Pont (Cotentin, Manche). x 20.
Voigt Collection, Hamburg, Nr. 3908.
FIG. 2. Young worn zoarium. From the matrix of D . 181. Upper Maastrichtian (Cotentin,
Manche). X2o. D. 49723.
Beisselina striata (Goldfuss) (p. 39)
FIG. 3. Worn fragment. From the matrix of D.iSi. Upper Maastrichtian (Cotentin,
Manche). x 20. 0.49721.
FIG. 4. Worn fragment for comparison with fig. 3. Upper Maastrichtian, Md., Geulem
near Berg, Geul-valley near Maastricht (Netherlands). Presented by the author, x 20.
D. 49722.
Frurionella fer tilts n. sp. (p. 38)
FIG. 5. Holotype, well preserved fragment with ovicelled zooecia. Upper Maastrichtian,
Chef du Pont (Cotentin, Manche). Presented by the author, x 20. 0.49610.
FIG. 6. Fragment showing zooecia with damaged ovicells. Upper Maastrichtian, Chef du
Pont (Cotentin, Manche). Presented by the author. X2O. 0.49612.
FIG. 7. Worn fragment, similar to fig. 6. From the matrix of D . 181. Upper Maastrichtian
(Cotentin, Manche). x 20. 0.49611.
Frurionella europaea (Voigt) (p. 39)
FIG. 8. For comparison with Frurionella fertilis n. sp. Upper Maastrichtian, Chef du Pont
(Cotentin, Manche). Presented by the author, x 20. 0.49720.
Vincularia canalifera (v. Hagenow) (p 27) (See also pi. 5, figs. 7-10)
FIG. 9. Small fragment. From the matrix of D.iSi. Upper Maastrichtian (Cotentin,
Manche). x 20. 0.49735.
FIG. 10. Badly preserved branched fragment showing an avicularium in the middle of the
right side. From the matrix of D.iSi. Upper Maastrichtian (Cotentin, Manche). x 20.
0.49736.
FIG. 1 1 . Fragment showing an avicularium at the right upper corner. Upper Maastrichtian
Md., Geulem near Berg, Geul-valley near Maastricht (Netherlands) . x 20. D . 49842. E. Voigt
Collection.
FIG. 12. Part of a branched stem, for comparison with fig. 9, showing relatively small zooecia
and opesia. Upper Maastrichtian, locality as fig. n. 0.49732.
Vincularia concinna (d'Orbigny) (p. 28)
FIG. 13. Holotype. Branched fragment for comparison with Vincularia canalifera von
Hagenow. Upper Maastrichtian, Nehou (Cotentin, Manche). x 20. d'Orbigny Collection,
Paris, Musee d'Histoire Naturelle, Nr. 7756.
Vincularia flexuosa (d'Orbigny) (p. 28)
FIG. 14. Holotype. Fragment for comparison with Vincularia canalifera von Hagenow.
Labelled as Santonian Venddme, Loir et Cher, France, x 20. d'Orbigny Collection, Paris,
Mus6e d'Histoire Naturelle, Nr. 7752.
Bull. BY. Mus. nat. Hist. (Geol.) 17, i
PLATES
PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING
THE CAUDAL SKELETON IN
MESOZOIC ACANTHOPTERYGIAN
FISHES
COLIN PATTERSON
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 2
LONDON: 1968
THE CAUDAL SKELETON IN MESOZOIC
ACANTHOPTERYGIAN FISHES
»
BY
COLIN PATTERSON
British Museum (Natural History)
Pp. 47-102; 28 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 2
LONDON: 1968
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
Papers was instituted, numbered serially for each
Department.
This paper is Vol. 17, No. 2 of the Geological
(Palaeontological) series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation:
Bull. Br. Mus. nat. Hist. (Geol.)
Trustees of the British Museum (Natural History) 1968
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 19 November, 1968 Price
THE CAUDAL SKELETON IN MESOZOIC
ACANTHOPTERYGIAN FISHES
By COLIN PATTERSON
CONTENTS
Page
I. INTRODUCTION ......... 49
II. SYSTEMATIC DESCRIPTIONS ....... 50
Order Ctenothrissiformes ....... 50
Order Beryciformes . . . . . . . . 56
Suborder Polymixioidei . . . . . . 56
Suborder Dinopterygoidei ...... 62
Suborder Berycoidei ....... 67
Order Lampridiformes ....... 78
Order Salmoniformes, Suborder Myctophoidei . . . 81
III. DISCUSSION .......... 83
(a) The relationships of Ctenothrissiformes, Myctophoidei and
Beryciformes ........ 83
(b) The origin of Perciformes ...... 87
(c) Intermediate groups ....... 96
IV. CONCLUSIONS .......... 98
V. REFERENCES .......... 100
VI. ABBREVIATIONS USED IN FIGURES ...... 102
SYNOPSIS
This paper contains descriptions of the caudal skeleton in all Ctenothrissiformes and Mesozoic
acanthopterygians, comparisons with living relatives being made where possible, and a brief
account of the caudal skeleton in Mesozoic Myctophoidei. These groups have a basically similar
caudal skeleton with the first ural and pre-ural centra fused, the second ural centrum free, six
hypurals, three epurals, a stegural and a second uroneural. Caudal scutes persist in Ctenothris-
siformes and primitive Myctophoidei and there is a single urodermal in some Cretaceous
myctophoids.
The bearing of caudal structure on the origin and early evolution of acanthopterygians is
discussed and variations in the second pre-ural neural spine are shown to be important. This
structure is primitively short and slender: two opposite developments from this condition are
elongation of the spine to support procurrent fin-rays (as in Polymixioidei) and reduction to a
low crest (as in Berycoidei and Perciformes). Wherever a full second pre-ural neural spine
occurs in perciform and higher groups it is produced secondarily by fusion with the first epural.
The evidence of the caudal skeleton does not support a polyphyletic origin of Perciformes from
different groups of Beryciformes. The Berycoidei appear to be the beryciform group most
closely related to the Perciformes.
The Danian Bathysoma is shown to be a lampridiform : the lampridiforms probably arose from
near Aipichthys and Pharmacichthys. The Zeiform.es are close relatives of the Berycoidei.
I. INTRODUCTION
IN an earlier paper (Patterson 1964) I gave detailed accounts of the skeletal
anatomy of the known Mesozoic acanthopterygian fishes (see also Patterson 1967)
but without describing the caudal skeleton. This was omitted because when the bulk
GEOL. 17, 2 5
50 THE CAUDAL SKELETON IN
of the work was done there existed no comprehensive description or even terminology
of the caudal skeleton of living teleosts. But during the last few years, following
from the work of Gosline (1960), the caudal skeleton has become recognized as an
important structure in tracing the relationships of teleost fishes and we now have
both a satisfactory terminology (Gosline 1960; Nybelin 1963; Monod 1967;
Patterson 1968) and a reasonable idea of the composition of the caudal skeleton
in most teleostean groups (Gosline 1960, 1961, ig6ia, 1963, 1965; Norden 1961;
Rosen 1962, 1964; Greenwood, Rosen, Weitzman & Myers 1966; Monod 1967;
Greenwood 1967; Weitzman 1967). In the light of this new information it seemed
that an investigation of the caudal skeleton in Mesozoic acanthopterygians might
prove a useful check on hypotheses of the origins of acanthopterygians and par-
acanthopterygians (Greenwood et al. 1966) . Also included are accounts of caudal
structure in the Ctenothrissiformes and, more briefly, in the Mesozoic Myctophoidei.
The terminology used here is that of Nybelin (1963) with the distinction between
" urodermal " and " uroneural " introduced by Patterson (1968) and the addition
from Monod (1967) of the terms " parhypural ", for the haemal arch of the first
pre-ural centrum, the terminal structure perforated by the caudal vein and artery
and which bears the hypurapophysis (Nursall 1963), and " stegural " for the paired
structure which articulates with the dorso-lateral surface of the first pre-ural centrum
and has a slender shaft extending postero-dorsally, lateral to the nerve cord. Monod
considers the stegural to be an element sui generis, not a compound structure.
In my opinion (Patterson 1968), the stegural represents the first uroneural fused
with neural arch material from the first pre-ural and first ural vertebrae. The term
stegural is nevertheless useful, since it obviates repetition of the cumbersome term
" first uroneural fused with the neural arches of the first ural and pre-ural centra ".
The material used is mainly in the collections of the British Museum (Natural
History) : these specimens are referred to by a registered number without prefix
or with the prefix " P ". Specimens from the American University, Beirut, the
American Museum of Natural History, New York, the Royal Scottish Museum,
Edinburgh, and the United States National Museum, Washington, are referred to
with the prefixes " AUB ", " AMNH ", " RSM " and " USNM " respectively. The
illustrations of fossil species are not reconstructions but camera lucida drawings of
single specimens.
I am grateful to Drs. P. H. Greenwood and D. E. Rosen, who have read and
criticized parts of this paper in manuscript, and for the loan of specimens to Prof.
T. Raven, American University, Beirut, Drs. C. D. Waterston and R. S. Miles, Royal
Scottish Museum, Dr. Bobb Schaeffer, American Museum of Natural History, and
Drs. D. H. Dunkle and D. M. Cohen, United States National Museum.
II. SYSTEMATIC DESCRIPTIONS
Order CTENOTHRISSIFORMES (Patterson 1964 : 218)
According to Marshall (1961) the living Macristium chavesi is a surviving cteno-
thrissiform (see also Greenwood, Rosen, Weitzman & Myers 1966; Patterson 1967 :
86). Unfortunately, the only extant specimen of Macristium is a post-larval
MESOZOIC ACANTHOPTERYGIAN FISHES 51
individual and the caudal skeleton is not ossified. But there appear to be six hypurals,
as in Ctenothrissiformes. Berry & Robins (1967) have described a second macristiid,
Macristiella perlucens, known only by a single post-larval specimen. In this fish
there are six hypurals, as in Ctenothrissiformes, but Berry & Robins think it unlikely
that the Macristiidae are close to the Ctenothrissiformes.
Family AULOLEPIDIDAE Patterson (1964 : 247)
Genus PATEROPERCA Smith Woodward (1942 : 543)
The type and only species, P. libanica Smith Woodward from the Middle Ceno-
manian of Hajula, Lebanon, is known only by two specimens, AUB 108906 (the
holotype) and AUB 108904. In 108904 the caudal region is badly preserved and
yields no useful information, but in the holotype the caudal skeleton is very well
preserved (Fig. i). The second pre-ural centrum (pu2] has a fully developed neural
spine (npu2) and an autogenous haemal arch. It is not possible to see with certainty
whether the haemal arch of the third pre-ural centrum (Pu3) is autogenous. The
first pre-ural and first ural centra are fused (pu i + ui) , these fused centra bearing
the parhypural (ph) and the first and second hypurals (hi, h2) in the normal way.
The first hypural is rather slender, only a little broader than the parhypural and
u2
npu2
pu1+u1
FIG. i. Pateroperca libanica Smith Woodward. Caudal skeleton of the holotype, AUB
108906, standard length 86 mm., Middle Cenomanian, Hajula, Lebanon. For explanation
of lettering see p. 102. Arrows mark the outermost (unbranched) principal fin-rays.
52 THE CAUDAL SKELETON IN
the second. Dorsally, the fused first pre-ural and ural centra bear a large stegural
(st) and there is a second uroneural (d2) ending, as usual, in front of the base of the
first unbranched principal ray of the caudal fin. There are three epurals (ei — j),
the third small and short. There is a small independent second ural centrum (u2)
bearing three hypurals (hj — 5) decreasing in size upwards, and above these there
is a small sixth hypural (h6) which probably failed to make contact with the ural
centrum. The foremost procurrent rays of the caudal fin, which are unsegmented
lepidotrichia, not spines, are inserted in front of the neural and haemal spine of the
third pre-ural centrum. The uppermost principal ray of the fin (unbranched) is
preceded by nine rays, only the last two or three segmented, and the lowermost by
seven rays, the last two segmented.
FIG. 2. Aulolepis typus Agassiz. Caudal skeleton of 47932, standard length c. 165 mm.,
Lower Chalk, Burham, Kent. /. r, foremost procurrent fin-ray (displaced) ; for explana-
tion of other lettering see p. 102. The upper hypurals are displaced ventrally, the
epurals and second uroneural are missing.
MESOZOIC ACANTHOPTERYGIAN FISHES
53
Genus AULOLEPIS Agassiz (1844 : 109)
In the type and only species, A. typus Agassiz from the Upper Cenomanian of
the English Chalk, only two specimens show any details of the caudal skeleton,
4033 (the holotype, Fig. 3) and 47932 (Fig. 2), and in both the bones are disturbed
to some extent. The caudal skeleton of Aulolepis agrees with that of Pateroperca
in most respects : fusion of the first pre-ural and first ural centra (pui + ui) , presence
of a free second ural centrum (u2], three epurals (ei — 3), six hypurals (hi — 6)
of which the first and the third are largest, an autogenous haemal arch on the second
pre-ural centrum (pu2), a stegural (st) and a free second uroneural (d2). The haemal
arch of the third pre-ural centrum (pu^] appears to be autogenous in 47932 but fused
with the centrum in 4033. Aulolepis differs clearly from Pateroperca in having the
neural spine of the second pre-ural centrum developed as a low, broad crest (npu2)
whose hind edge fits between the front edges of the stegurals. In my earlier des-
criptions of Aulolepis and Pateroperca (Patterson 1964 : 247) I discussed the difficulty
of separating the two genera and considered the possibility that P. libanica is merely
npu2
d2
pu1+u1
FIG. 3. Aulolepis typus Agassiz. Caudal skeleton of the holotype, 4033, standard length
c. 135 mm., Lower Chalk, Lewes, Sussex. For explanation of lettering see p. 102. The
epurals and upper hypurals are displaced ventrally, the stegural is displaced forwards.
54
THE CAUDAL SKELETON IN
a species of Aulolepis, but the presence on the second pre-ural centrum of a fully
developed neural spine in Pateroperca and of a low crest in Aulolepis clearly separates
the two genera. The caudal of 47932 shows one other point of interest, a slender,
elongated caudal scute (/. s, Fig. 2) in front of the upper lobe of the fin. There is
no conclusive evidence of a similar structure in front of the lower lobe of the fin,
but it was probably present since there is a caudal scute in front of each lobe of the
fin in Ctenothrissa (see below) and in most living teleosts which still retain these
structures (Elops, Tarpon, Albula, Aulopus and Chanos, Gosline 1965 : 192; Argen-
tina, Weitzman 1967 : 532). There is no sign of caudal scutes in either specimen of
Pateroperca, but again it is probable that they were present. No specimen of Aulol-
epis has the fin-rays of the caudal fin well preserved but 4033 shows that the foremost
procurrent rays of the lower lobe articulated with the haemal spine of the third
pre-ural centrum, and both 4033 and 47932 show that these foremost rays were true
spines.
Family CTENOTHRISSIDAE Smith Woodward (1901)
Genus CTENOTHRISSA Smith Woodward (1899 : 490)
In Ctenothrissa, the only genus of the Ctenothrissidae, the caudal skeleton and fin
are well exposed in the two species from the Cenomanian of the Lebanon, C. vexillifer
e3
u2
pu1+u1
FIG. 4. Ctenothrissa signifer Hay. Caudal skeleton of P. 47524, standard length 83 mm.,
Middle Cenomanian, Hajula, Lebanon. For explanation of lettering see p. 102. Arrows
mark the outermost (unbranched) principal fin-rays.
MESOZOIC ACANTHOPTERYGIAN FISHES
55
(Pictet 1850), the type species, and C. signifer Hay (1903) (Fig. 4), and the caudal
skeleton is preserved in one or two specimens of C. microcephala (Agassiz) (Fig. 5)
and C. radians (Agassiz) from the Upper Cenomanian of the English Chalk. There
seem to be no significant differences in caudal anatomy between these four species.
As in Aulolepididae, there is a free second ural centrum (u2), the first ural and first
pre-ural centra are fused (pui + ui], there are six hypurals (hi — 6), the first and
third the largest and the sixth failing to articulate with the second ural centrum,
there are three epurals (ei — j), a stegural (st) and a second uroneural (£2) and the
haemal arch of the second pre-ural centrum (pu2) is autogenous. The haemal arch
of the third pre-ural centrum (pu>3) appears partially or completely fused to the
centrum. As in Aulolepis, but in contrast to Pateroperca, the neural spine of the
second pre-ural centrum (npu2) is represented by a broad crest, about half as high
as the preceding spine. There is a slender caudal scute (/. s.) in front of both the
upper and lower lobes of the caudal fin. In C. vexillifer and C. signifer, the only
species in which the caudal fin-rays are well preserved, both the upper and lower
principal rays are preceded by six rays, the last two segmented, and the foremost
fin-rays are inserted on the neural and haemal spines of the third pre-ural centrum.
e3
e1
npu2
2mm
FIG. 5. Ctenothrissa microcephala (Agassiz). Caudal skeleton of 49881, standard length
c. 1 15 mm., Lower Chalk, zone of Holaster subglobosus, Reigate, Surrey. For explanation
of lettering see p. 102. The upper hypurals are displaced ventrally. Of the two elements
labelled ? h6, that on the left may be the base of a fin-ray.
56 THE CAUDAL SKELETON IN
In these small species from the Lebanon, the foremost procurrent caudal rays are
unsegmented lepidotrichia, with separate right and left halves, but in the larger
species from the English Chalk they appear to be true spines.
Order BERYCIFORMES
Suborder POLYMIXIOIDEI Patterson (1964 : 433)
Family POLYMIXIIDAE Gill (1862)
The only living genus of this family and suborder is Polymixia. The caudal
skeleton of Polymixia nobilis has been figured by Regan (1911, fig. i) and briefly
discussed by Gosline (1961 : 14). The specimen illustrated here (Fig. 6) is almost
certainly that used by Regan. The haemal arches of the second and third pre-ural
centra (pu2, pus) are autogenous, the second pre-ural centrum has a fully developed
neural spine (npu2) , the first pre-ural and first ural centra are fused (pui + ui) and
there is a free second ural centrum (14,2). There is a large stegural (st) and a free
second uroneural (d2). There are three slender epurals (ei — j) and six hypurals
(hi — 6) of which the first and the fourth are the largest, the third being excavated
posteroventrally to give a notch between the hypurals supporting the upper and
lower lobes of the fin. The second ural centrum has a long posterior process and
makes contact with the sixth hypural. The first rays of the caudal fin articulate
with the neural and haemal spines of the third pre-ural centrum. In the upper lobe
of the fin the first unbranched principal ray is preceded by four true spines and two
e1-3
npu2
pu1+u1
u2
FIG.
3mm
6. Polymixia nobilis Lowe. Caudal skeleton of a dried skeleton, 1895.5.28.1,
standard length 220 mm., Madeira. For explanation of lettering see p. 102.
MESOZOIC ACANTHOPTERYGIAN FISHES
segmented rays, in the lower lobe by three spines and two segmented rays,
caudal skeleton of P. japonicus does not differ from that of P. nobilis.
57
Genus BERYCOPSIS Dixon (1850 : 372)
In the type species, B. elegans Dixon from the Upper Cenomanian and Turonian
of the English Chalk, the caudal skeleton is well preserved in 25881 (Fig. 7) and
P. 6465. The caudal skeleton of B. elegans agrees with that of Polymixia in almost
every detail except that the third hypural (Aj) is not excavated postero-ventrally
so that it is as large as the fourth, the haemal arch of the third pre-ural centrum,
probably autogenous in P. 6465, is fused with the centrum in 25881, and the third
and fourth hypurals, separate in 25881 (standard length c. 13 cm.), are completely
fused in the larger P. 6465 (standard length c. 16 cm.). In 25881 the lateral surface
of the second ural centrum is covered by a small, triangular plate (x, Fig. 7).
P. 6465 is not sufficiently well preserved to see whether this plate is present. The
plate appears to be part of the caudal skeleton, not a dermal element which has
become apposed to the centrum post mortem. The only record of a similar structure
is in the living myctophoid Synodus foetens, where Hollister (193 ja, figs. 5, 12) found
a " uroneural " in this position, ventral to the normal uroneurals. The foremost
h6
h5
npu2
pul+ul
FIG. 7. Berycopsis elegans Dixon. Caudal skeleton of 25881, standard length c. 130 mm.,
Chalk, Sussex, x, unidentified plate covering second ural centrum; for explanation of
other lettering see p. 102.
58 THE CAUDAL SKELETON IN
unsegmented caudal fin rays of B. elegans are soft rays, not spines as they are in
Polymixia.
Of P. 1047 1, a small fish lacking the head, I rashly stated that it is " certainly
B. elegans " (Patterson 1964 : 278), but on re-examining this specimen I find that the
scales are larger than in B. elegans, the dorsal and anal spines are longer, there are
five anal spines (not four as stated earlier) and in the caudal skeleton the neural
spine of the second pre-ural centrum is much reduced. This specimen is clearly not
B. elegans and the reduced neural spine of the second pre-ural centrum shows that
it is not even a polymixioid, but there is insufficient evidence to decide on its real
position.
The second species of Berycopsis, B. germanus (Agassiz) from the Campanian of
Westphalia, shows nothing in the caudal skeleton to distinguish it from B. elegans
except that the haemal arch of the third pre-ural centrum is clearly autogenous and
there is no sign of fusion between the third and fourth hypurals in the two specimens
where this region is visible. B. germanus is so preserved that it shows the details
of the caudal fin-rays much better than B. elegans. The foremost rays of the caudal
fin are arranged asymmetrically, the upper rays articulating with the neural spine
of the second pre-ural centrum, the lower with the haemal spine of the third pre-ural
centrum. The first principal ray (unbranched) of the upper lobe is preceded by
four unsegmented and one segmented ray, the lower by three unsegmented and one
segmented ray. It is not possible to see whether the foremost rays are spines (as
in Polymixia) or soft rays, as in B. elegans and Omosoma.
Genus OMOSOMA Costa (1857 : Io6)
Having now had the opportunity to make a direct comparison between the holo-
types of Omosoma pulchellum (Davis 1887) (RSM 1891.59.72) and 0. intermedium
Smith Woodward (1901) (48112) I find that the two are conspecific, the median fin
counts (D V, 28-30; A III-IV, 24) being the same in both. The apparent differences
in proportions of the two (Smith Woodward 1901 : 420) are due to longitudinal
compression of the holotype of 0. pulchellum, the holotype of 0. intermedium showing
the true proportions of the fish. In my earlier description of Omosoma (Patterson
1964 : 374) the median fin counts given for 0. pulchellum (D IV-V, 35 ; A III-IV, 29)
were based on distorted specimens of 0. sahelalmae. 0. intermedium is therefore a
synonym of 0. pulchellum.
None of the specimens of Omosoma available, belonging to 0. sahelalmae Costa
(the type species) and 0. pulchellum (Davis), from the Upper Santonian of Sahel
Alma, Lebanon (Ejel & Dubertret 1966, have recently found evidence of the precise
age of these beds), has the caudal skeleton sufficiently well preserved to be illustrated,
but all give a picture of a caudal skeleton which does not differ significantly from that
of Berycopsis. The foremost rays of the caudal fin are arranged asymmetrically,
those of the upper lobe articulating with the neural spine of the second pre-ural
centrum, those of the lower lobe with the haemal spine of the third pre-ural centrum,
just as in B. germanus. In both the lobes of the fin there are 4 unsegmented
lepidotrichia (not spines) and two segmented rays in front of the principal rays.
MESOZOIC ACANTHOPTERYGIAN FISHES
59
It is impossible to see whether the haemal arch of the third pre-ural centrum is
autogenous.
Genus PYCNOSTERINX Heckel (1849 : 337)
Pycnosterinx is known by four species, all from the Upper Santonian of Sahel
Alma, Lebanon. Figure 8 shows the caudal skeleton of P. russeggerii Heckel, the
type species: it is very similar to those of the other polymixiids and differs from
Polymixia (Fig. 6) only in having the postero-ventral part of the third hypural com-
pletely ossified, so that there is no gap between the hypurals supporting the upper
and lower lobes of the fin. As in Polymixia the neural and haemal spines are strongly
inclined and the epurals slender. As in Berycopsis and Omosoma, but in contrast
to Polymixia, the foremost caudal fin-rays are arranged asymmetrically, the upper
ones articulating with the neural spine of the second pre-ural centrum, the lower
with the haemal spine of the third pre-ural centrum. In both the upper and lower
lobes of the fin the principal rays are preceded by three unsegmented soft rays and
three segmented rays. P. discoides Heckel and P. gracilis Davis do not differ in
caudal structure from P. russeggerii except that in P. gracilis there appear to be
only five procurrent rays. I have seen no specimens of P. dubius Davis in which
the caudal skeleton and fin are well preserved, but in the holotype of Pycnosterinx
latus Davis (1887 : 534, pi. 27, fig. 2), RSM 1891.59.77 (referred to as a "frag-
mentary fish of indeterminable genus " by Smith Woodward 1901 : 395), the scales
have the same spinous surface as in P. dubius (Patterson 1964 : 380) and it is probable
that the specimen is a large, distorted individual of P. dubius, although the state of
u2
pu1>u1
2mm
FIG. 8. Pycnosterinx russeggerii Heckel. Caudal skeleton of 47820, standard length
78 mm., Upper Santonian, Sahel Alma, Lebanon. For explanation of lettering see
p. 102.
6o
THE CAUDAL SKELETON IN
preservation of this and the holotype of P. dubius is such that this cannot be demon-
strated with sufficient certainty to synonymize the two species. P. latus appears
to have one or two more anal fin-rays and caudal vertebrae than the holotype of
P. dubius. The caudal skeleton is completely exposed in the holotype of P. latus
and does not differ from that of P. russeggerii in any way, but both in this specimen
and in P. dubius the foremost fin-rays are true spines, a difference from the other
species of Pycnosterinx and a resemblance to the living Polymixia.
Genus HOMONOTICHTHYS Whitley (1933 : 146)
All three species of this genus occur in the Upper Cenomanian — Lower Turonian
of the English Chalk. In the type species, H. dorsalis (Dixon), only one specimen,
43575, shows anything of the caudal skeleton and here it is only possible to see that
there was a fully developed neural spine on the second pre-ural centrum and that
the foremost rays of the upper caudal lobe are true spines, articulating with the
neural spine of the third pre-ural centrum. In these last two features H. dorsalis
e2
e3
d2
2mm
FIG. 9. Homonotichthys pulchellus (Dixon). Caudal skeleton of P. 11112, standard length
c. no mm., Lower Chalk, Amberley, Sussex. For explanation of lettering see p. 102.
MESOZOIC ACANTHOPTERYGIAN FISHES 61
resembles Polymixia and differs from other Cretaceous polymixiids except
Pycnosterinx dubius.
In H. pulchellus (Dixon) the caudal skeleton is preserved in the holotype, 25886,
and in two specimens only recently recognized as belonging to this species, P. 11112
(Fig. 9) and P. 10639: the latter are about no and 120 mm. respectively in standard
length, the largest examples of this species yet recorded. The caudal skeleton of
P.IIH2 (Fig. 9) is abnormal in the partial doubling of the neural spine of the second
pre-ural centrum and in having the second epural smaller than the third. The
caudal skeleton of H. pulchellus is very like that of Polymixia, with autogenous
haemal arches on the second and third pre-ural centra (pu2, pu3) six hypurals
(hi — 6), three slender epurals (ei — 3), a free second ural centrum (u2), a stegural
(st) and a second uroneural (d,2}. The postero- ventral corner of the third hypural
is truncated, but less so than in Polymixia. Broad flanges on the anterior margin
of the neural spines of the second and third pre-ural centra are characteristic of
H. pulchellus. All three specimens show that the foremost caudal rays are spines,
as in H. dorsalis and Polymixia. In P . 10639 the caudal rays articulate with the
neural and haemal spines of the third pre-ural centrum, as in Polymixia. In P . 11112
where the second pre-ural spine is abnormal, the foremost upper caudal rays articu-
late with the neural spine of the second pre-ural centrum. The lowermost principal
caudal ray (unbranched) is preceded by four spines and three segmented rays.
P.iiii2 shows the anal fin of H. pulchellus, previously unknown. The fin contains
five spines, the first very small and the fifth the longest and thickest, and about
eleven soft rays. The fifth anal spine is equal in length to the longest dorsal spine,
just over one-quarter of the maximum depth of the trunk.
H. rotundus (Smith Woodward) is known only by the holotype, P . 315, and P . 5682.
P . 5682 shows most of the details of the caudal skeleton : there is nothing to distin-
guish it from the other polymixiids described here except that there is no flange
on the anterior margin of the neural spine of the second pre-ural centrum as there
is in H. pulchellus. P. 315 shows that the foremost caudal rays are soft rays,
longitudinally divided, not spines as they are in H. dorsalis, H. pulchellus and Poly-
mixia. These two points may be added to others (Patterson 1964 : 299) indicating
that this species is incorrectly placed in Homonotichthys, but more specimens are
necessary before its true position can be decided.
Family SPHENOCEPHALIDAE Patterson (1964 : 383)
The only member of this family is Sphenocephalus fissicaudus Agassiz from the
Campanian of Westphalia. Of the six specimens in the British Museum (Natural
History), three, P. 2100 (Fig. 10), P. 8772 and P. 9059 have the caudal skeleton well
preserved. As in Polymixiidae, the second pre-ural centrum has a fully developed
neural spine (npu2], there is a free second ural centrum (1*2), a stegural (st), a second
uroneural (d2), and six hypurals (hi — 6). As in Polymixia, the foremost procurrent
rays articulate with the neural and haemal spines of the third pre-ural centrum.
In contrast to all Polymixiidae there are only two epurals (ei, 2), the first curved
forwards proximally and with a gap between it and the neural spine of the second
62
THE CAUDAL SKELETON IN
FIG. 10. Sphenocephalus fissicaudus Agassiz. Caudal skeleton of P. 2100, standard length
92 mm., Campanian, Sendenhorst, Westphalia. For explanation of lettering see p. 102.
pre-ural centrum, there is a wide gap between the upper and lower hypurals, and
there is a large number of procurrent rays, apparently nine in each lobe, six
unsegmented soft rays and three segmented in the upper, five unsegmented and
four segmented in the lower.
Suborder DINOPTERYGOIDEI Patterson (1964 : 434)
This suborder contains four monotypic Uppper Cretaceous familes, probably not
closely related. As the caudal skeleton in the type family, the Dinopterygidae, is
poorly known, the best known family, the Aipichthyidae, will be described first.
Family AIPICHTHYIDAE Patterson (1964 : 303)
Genus AIPICHTHYS Steindachner (1860 : 763)
I have seen no material of the type species, A. pretiosus Steindachner from the
Lower Cenomanian of Dalmatia, and the caudal region is not preserved in A.
MESOZOIC ACANTHOPTERYGIAN FISHES 63
nuchalis (Dixon) from the English Chalk. The other two species, A. minor (Pictet)
and A. velifer Smith Woodward, both from the Cenomanian of Hakel, Lebanon,
are well represented in the British Museum (Natural History). They show a
remarkable range of variation in the structure of the caudal skeleton. In most speci-
mens of Aipichthys the distal parts of the hypurals, epurals and uroneurals are
obscured by the deeply cleft bases of the caudal fin-rays, but in P. 82 (A. minor,
Fig. uA) the fin-rays are displaced, exposing these bones. In this specimen and in
all others the neural and haemal spines of the third pre-ural centrum (pu3) are broad
and elongate and the first procurrent rays of the fin articulate with them. The
haemal arches of the second and third pre-ural centra are autogenous. In contrast
to all Polymixioidei, the neural spine of the second pre-ural centrum (npu2) is
normally only about half as long as its predecessor, which makes contact with the
first epural distally. Though somewhat expanded, this neural spine is more like
those of Elops (Nybelin 1963, figs, i, 4) and Nematonotus (Fig. 25) than the shorter,
broader spine in Aulolepis and Ctenothrissa (Figs. 2, 4, 5) : it does not resemble the
very low crest on this centrum in Berycoidei and percoids. In one specimen of
Aipichthys minor, P. 6183 (Fig. nC), the neural spine of the second pre-ural centrum
is fully developed and supports epaxial fin-rays, as in Polymixioidei. This is
clearly an individual variation, comparable to those found in certain individuals of
Monocentris (Fig. 14) and Siniperca (Fig. 26), and is recognizable as such by the gap
between the spine and the proximal part of the first epural, which is filled by flanges
from the posterior face of the spine and from the anterior face of the epural. In
all specimens of Aipichthys there are three epurals (ei — 3) and the first ural and
pre-ural centra are fused (pui + ui). There is normally a free second ural centrum
(u2, Figs. 1 1 A, C), but in occasional individuals, like the large specimen shown in
Fig. nB, the second ural centrum is fused into the preceding compound centrum
(pui + ui + U2), although the line of fusion is visible in transparency under xylene.
There is always a stegural (st) and a second uroneural (d2). In A. minor there are
normally six hypurals (hi — 6, Figs. nB, C), as in Polymixioidei, but in P. 82 (Fig.
1 1 A) the fifth hypural is partially divided distally suggesting that the sixth is
fused into it. In all specimens of A. velifer in which the hypurals are visible (P. 4743,
P. 4744, 49486, P. 47862) there appear to be only five hypurals. Preceding the
principal rays of the caudal fin there are four unsegmented soft rays and three seg-
mented rays in the upper lobe, three unsegmented and three segmented rays in the
lower.
Family PHARMAGICHTHYIDAE Patterson (1964 : 398)
This family contains only Pharmacichthys venenifer Smith Woodward (1942) from
the Middle Cenomanian of Hakel, Lebanon. None of the five known specimens
of this species has the caudal skeleton sufficiently well preserved to be illustrated,
but the holotype, AUB 104691/99, and AUB 101872 show most of the caudal anatomy.
The suggestion (Patterson 1964 : 401) that the nearest relative of Pharmacichthys
is Aipichthys is borne out by the structure of the caudal skeleton and fin. In almost
every respect the caudal skeleton of Pharmacichthys is identical with that of Aipi-
chthys (Fig. n). The foremost caudal fin-rays articulate with the neural and haemal
GEOL. I 7, 2 6
64
THE CAUDAL SKELETON IN
,e1-3
d2
npu2
h5
•pu1+u1+u2
MESOZOIC ACANTHOPTERYGIAN FISHES
e1-3
d2.
h6
puUul
u2
FIG. ii. Aipichthys minor (Pictet). Caudal skeleton of A, P. 82, standard length 32 mm.;
B, RSM 1881 .5.41, standard length c. 50 mm.; C, P. 6183, standard length 36 mm. All
from Middle Cenomanian, Hakel, Lebanon. For explanation of lettering see p. 102.
In B arrows mark the outermost (unbranched) principal fin-rays.
spines of the third pre-ural centrum, the neural spine of the second pre-ural centrum
is about half as long as its predecessor, as in Aipichthys, the haemal spine of the
second pre-ural centrum has a broad flange on its anterior edge, there are three
epurals and the principal rays of the fin are preceded by four unsegmented soft rays
and three segmented rays above, three unsegmented and three segmented rays below.
It is impossible to see whether there are five or six hypurals. A further resemblance
to Aipichthys, not previously noticed, is that the bases of the caudal rays are deeply
cleft, overlapping much of the hypurals (" hypurostegy ", Le Danois & Le Danois
1964). The only difference from Aipichthys, suggested by the holotype, AUB
101872 and AUB 102601, is that the first and second hypurals appear to be fused.
Family PYGNOSTEROIDIDAE Patterson (1964 : 389)
The only member of this family is Pycnosteroides levispinosus (Hay 1903) from the
Middle Cenomanian of Hajula, Lebanon. The caudal skeleton and fin are present
in two specimens, P. 13900 (Fig. 12) and AMNH 45190 (Hay 1903, pi. 32, fig. 3).
Pycnosteroides differs from Aipichthys and Pharmacichthys in having the foremost
rays of the caudal fin inserted on the first epural above and the haemal spine of
the third pre-ural centrum below, and in having a fully developed neural spine on
66
THE CAUDAL SKELETON IN
el
npu2,
st
FIG. 12. Pycnosteroides levispinosus (Hay). Caudal skeleton of P. 13900, standard
length c. 50 mm., Middle Cenomanian, Hajula, Lebanon. For explanation of lettering
seep. 1 02.
the second pre-ural centrum (npu2). This neural spine differs from those of the
Polymixioidei in being expanded distally. The haemal arches of the second and
third pre-ural centra (pu2, puj) are autogenous. In the fusion of the first ural and
first pre-ural centra (pui -f- ui), the free second ural centrum (u2), and the form of
the stegural (st} and second uroneural (missing in the figured specimen but present
in AMNH 45190;) Pycnosteroides agrees with Aipichthys and the Polymixioidei. There
are three epurals (ei — 3). In P. 13900, as in Aipichthys velifer, there are only five
hypurals (hi — 5), but here the first and third are the largest, the third being much
larger than the fourth. In AMNH 45190 there are six hypurals. In contrast to
Aipichthys and Pharmacichthys there are only three soft rays, all unsegmented, in
front of both the upper and lower principal rays, and the fin-rays are not deeply
cleft basally.
Family DINOPTERYGIDAE Jordan (1923 : 173)
This family, the type of the suborder, contains only Dinopteryx spinosus (Davis)
from the Upper Santonian of Sahel Alma, Lebanon.
The caudal region is very imperfectly preserved in the holotype,
MESOZOIC ACANTHOPTERYGIAN FISHES
67
but is more or less completely visible in USNM 22217 and 22219. As in other respects
(Patterson 1964 : 392), Dinopteryx resembles Pycnosteroides in the caudal skeleton.
The second pre-ural centrum has a fully developed neural spine and both this and
the preceding neural spine are expanded distally, as in Pycnosteroides (Fig. 12). As
in Pycnosteroides there are three epurals, the first ural and pre-ural centra are fused,
there is a free second ural centrum, a stegural and a second uroneural. USNM 22219
shows that there were at least three upper hypurals, shaped as in Pycnosteroides,
but it is impossible to be certain whether or not a small sixth hypural was present.
The foremost procurrent caudal rays articulate with the neural spine of the second
pre-ural centrum above and the haemal spine of the third pre-ural centrum below,
further forwards than in Pycnosteroides. There are four spines and two segmented
rays in front of the principal rays above, three spines and two segmented rays below,
both the holotype and USNM 22217 showing that the foremost procurrent rays were
true spines, a difference from other Dinopterygoidei.
Suborder BERYCOIDEI Patterson (1964 : 433)
This suborder, containing eight living families, is represented in the Cretaceous
only by two families, the Trachichthyidae and Holocentridae. A detailed discussion
of the relationships between these two families in the Cretaceous will be found in
npu3
pu1+u1
npu2
FIG. 13. Hoplostethus mediterraneus Cuvier & Valenciennes. Caudal skeleton of a dried
skeleton, 1878.4.5.8, standard length 190 mm., Japan. For explanation of lettering
seep. 102.
68 THE CAUDAL SKELETON IN
Patterson (1967). Among living Berycoidei, the caudal skeleton of Centroberyx
affinis (Berycidae) has been illustrated by Regan (1911, fig. 2), those of Hoplostethus
(Trachichthyidae), Holocentrus and Myripristis (both Holocentridae) are briefly
discussed by Gosline (1961 : 14) and a caudal skeleton of Holocentrus ascensionis is
figured by Rosen (1964, fig. 23D). Fig. 13 shows the caudal skeleton of the living
Hoplostethus mediterraneus (Trachichthyidae). In most respects this is typical of
primitive Berycoidei, with autogenous haemal spines on the second and third
pre-ural centra, the second pre-ural centrum without a neural spine, bearing only a
low, broad crest (npu2) which lies below the proximal end of the first epural, three
epurals (ei — 3), six hypurals (hi — 6), a free second ural centrum (u2), a stegural
(st) and a second uroneural (d2) . In the specimen illustrated the neural spine of the
third pre-ural centrum (npuj) is double distally and the third and fourth hypurals
are fused proximally: these features are individual abnormalities. The neural crest
on the second pre-ural centrum is autogenous — this is a primitive feature which also
occurs in primitive myctophoids (N ematonotus and Aulopus, p. 81). In Hoplo-
stethus the foremost procurrent fin-rays articulate with the neural and haemal spines
of the third pre-ural centrum. In living trachichthyids (Hoplostethus, Trachichthys,
Paratrachichthys, Gephyroberyx) the nineteen principal caudal rays are normally
preceded by six spines and one segmented ray above and below.
The caudal skeleton in living holocentrids is described below (p. 75, Fig. 20).
In Berycidae (Regan 1911, fig. 2) the caudal skeleton is advanced over the trachi-
chthyid condition in having both the second ural centrum and the stegural fused
into the compound first ural and pre-ural centrum. Nothing is known of the caudal
skeleton in Korsogasteridae and Anomalopidae. Dissection of single specimens of
Diretmus (Diretmidae) and Anoplogaster (Anoplogasteridae) shows that both
resemble Berycidae in having the stegural and second ural centrum fused into the
preceding compound centrum, while in Anoplogaster the second uroneural is lost
and in Diretmus the sixth hypural is lost and there is fusion between the first and
second hypurals and between the third and fourth hypurals. In the Stephano-
berycoidei, which appear to be only specialized derivatives of the Berycoidei,
Gibberichthys (Gibberichthyidae) agrees with trachichthyids such as Hoplostethus in
the caudal skeleton but Melamphaes (Melamphaeidae) has both the stegural and
second ural centrum fused with the preceding centrum, a much reduced second
uroneural, only five hypurals and fusion within the upper and lower hypurals. All
living berycoids and stephanoberycoids seem to be characterized by the presence
of spinous procurrent caudal rays.
In Monocentris (Monocentridae) the caudal skeleton normally shows the same
major features as the figured specimen of Hoplostethus (even to the partial doubling of
the neural spine of the third pre-ural centrum), but in one of the available skeletons
(Fig. 14) there is a fully developed neural spine on the second pre-ural centrum
(npu2) and the neural spine of the third pre-ural centrum is single. There are three
epurals (ei~3), so that in this individual the neural spine on the second pre-ural
centrum has apparently developed instead of the normal doubling of the neural
spine of the preceding centrum. The caudal skeleton of this specimen resembles
those of polymixioids and the dinopterygoids Pycnosteroides and Dinopteryx, but
MESOZOIC ACANTHOPTERYGIAN FISHES
69
FIG. 14. Monocentris japonicus (Houttuyn). Caudal skeleton of a dried skeleton showing
a neural spine on the second pre-ural centrum, 1862. u . i .47, standard length 125 mm.,
Japan. For explanation of lettering see p. 102.
it is questionable whether this is significant. However, Monocentris seems to be
the only living berycoid in which the procurrent caudal rays are sometimes not spines
but unsegmented soft rays : of four specimens examined, one has no spines and one
has only one spine in front of the lower caudal lobe and none above. Monocentris
is a fish of highly specialized appearance and the skull suggests that the Monocent-
ridae are derivatives of the Trachichthyidae, but there is a patch of teeth on the
endopterygoid (Starks 1904 : 618), a character otherwise unknown in Berycoidei,
and this and the occasional absence of procurrent caudal spines suggest that the
Monocentridae may have had a long independent history.
Family TRACHICHTHYIDAE Bleeker (1859)
The caudal skeleton of the living Hoplostethus mediterraneus is described above
(Fig. 13).
Genus HOPLOPTERYX Agassiz (1838 : 4)
Hoplopteryx, with eight species ranging from the Middle Cenomanian to the Upper
Senonian, is the longest-ranging and largest genus of Cretaceous Trachichthyidae.
7o
THE CAUDAL SKELETON IN
h6
h5
pu1+u1
FIG. 15. Hoplopteryx lewesiensis (Mantell). Caudal skeleton of P. 19486, standard length
c. 175 mm., Chalk, Sussex. For explanation of lettering see p. 102.
In the type species, H. antiquus Agassiz from the Campanian of Westphalia, I have
seen no specimens in which the caudal skeleton is preserved. The best known
species is H. lewesiensis (Mantell) which ranges throughout the English Chalk:
several specimens show the caudal skeleton (Figs 15, 16) which is almost identical
with that of the living Hoplostethus (Fig. 13), with the neural arch of the second
pre-ural centrum (npu2) reduced (though not so much as in Hoplostethus}, the haemal
arches of the second and third pre-ural centra autogenous, three epurals (ei~3) of
which the first is especially large, a stegural (sf) and a second uroneural (d2), a free
second ural centrum (u2) and six hypurals (hi-6), the uppermost very small. The
neural arch of the second pre-ural centrum is not autogenous as it is in Hoplostethus.
The foremost caudal fin-rays are inserted on the first epural above and on the haemal
spine of the third pre-ural centrum below. There are only three spines and one
segmented ray in front of the upper principal rays and two spines and one segmented
ray in front of the lower.
The other species of Hoplopteryx in the English Chalk are H. simus Smith Wood-
ward, H. macr acanthus Patterson and H. gephyrognathus Patterson. The caudal
MESOZOIC ACANTHOPTERYGIAN FISHES
U2
2mm
FIG. 16. Hoplopteryx lewesiensis (Mantell). Caudal skeleton of P. 5421, standard length
c. 135 mm., Lower Chalk, Lewes, Sussex. For explanation of lettering see p. 102. The
second uroneural is displaced ventrally and the fifth and sixth hypurals are missing.
skeleton of H. simus is exposed in P. 11202, that of H. macmcanthus in P. 30186:
neither appears to differ from H. lewesiensis in any way. The caudal region is not
preserved in the two known specimens of H. gephyrognathus.
The earliest species of Hoplopteryx is H. lewisi (Davis) from the Middle Ceno-
manian of Hakel, Lebanon. In this species the caudal skeleton and fin are exposed
in P. 10709 and partially shown in the holotype, P. 4758. H. lewisi seems to agree
with H. lewesiensis in every detail, even to the insertion of the foremost upper fin
rays on the first epural, except that there are four spines and one segmented ray in
front of the upper principal rays, three spines and one segmented ray in front of the
lower.
The remaining two species of Hoplopteryx, H. syriacus (Pictet & Humbert) and
H. spinulosus Smith Woodward, are from the Upper Santonian of Sahel Alma,
Lebanon. In H. spinulosus I have seen no specimen in which the caudal skeleton
72
THE CAUDAL SKELETON IN
is preserved. In H. syriacus parts of the caudal skeleton are preserved in 49553
and they show nothing to distinguish the species from H. lewesiensis.
Genus LISSOBERYX Patterson (1967 : 73)
The type species, L. dayi (Smith Woodward 1942), is from the M. Cenomanian
of Hakel and Hajula, Lebanon. I have briefly described the caudal skeleton (Pat-
terson 1967 : 78) which is preserved in AUB 108926 (Fig. 17) and AUB 101997.
The caudal skeleton agrees well with those of Hoplostethus (Fig. 13) and Hoplopteryx
(Figs 15, 16). The neural spine of the second pre-ural centrum (npu2) is reduced
to about the same extent as in Hoplopteryx, there are three epurals (21-3), the
d2
st
npu2
pu3
FIG. 17. Lissoberyx dayi (Smith Woodward). Caudal skeleton of AUB 108926, standard
length 34 mm., Middle Cenomanian, Hajula, Lebanon. For explanation of lettering
see p. 102. Arrows mark the outermost (unbranched) principal fin-rays.
first long and closely applied distally to the neural spine of the third pre-ural centrum,
a stegural (st) and a second uroneural (d,2), and six hypurals (hi-6). The only
significant differences from Hoplopteryx and Hoplostethus are that the fused first ural
and pre-ural centrum is longer, with clear signs in the surface sculpture of its origin
from two centra, and that the second ural centrum is also longer, so that the caudal
skeleton appears more upturned. As in Hoplopteryx the foremost caudal rays
articulate with the first epural above and the haemal spine of the third pre-ural
centrum below. The principal rays are preceded by four spines and one unseg-
mented ray above, three spines and one segmented ray below.
MESOZOIC ACANTHOPTERYGIAN FISHES 73
Genus ACROGASTER Agassiz (1838 : 5)
I have seen no specimens of A. parvus Agassiz, the type species, or of A. brevi-
costatus von der Marck, both these species from the Campanian of Westphalia being
poorly known. The remaining species, A. heckeli (Pictet) and A. daviesi (Davis),
from the Upper Santonian of Sahel Alma, Lebanon, are common and several specimens
in the British Museum (Natural History) show the caudal skeleton clearly: I can
find no differences between these two species in caudal anatomy. A specimen of
u2
FIG. 18. Acrogaster heckeli (Pictet). Caudal skeleton of P. 4155, standard length c. 55 mm.
Upper Santonian, Sahel Alma, Lebanon. For explanation of lettering see p. 102.
A. heckeli is illustrated in Fig. 18. As in the other trachichthyids described above,
the neural spine of the second pre-ural centrum is reduced (npu2), there are three
epurals (ei-j), a stegural (st), a second uroneural (d2) and six hypurals (hi-6).
The fused first ural and pre-ural centra (pui + ui) and the second ural centrum (u2)
are elongate, as in Lissoberyx, and the caudal skeleton appears strongly upturned.
As in Hoplopteryx and Lissoberyx, the foremost caudal rays articulate with the first
epural above and the haemal spine of the third pre-ural centrum below. The
principal caudal rays are preceded by four or five spines and one segmented ray above,
three or four spines and one segmented ray below.
74
THE CAUDAL SKELETON IN
Genus TUBANTIA Patterson (1964 : 413)
The only species is T. cataphractus (von der Marck), from the Campanian of
Westphalia, in which the caudal skeleton is well exposed in P. 21984 (Fig. 19).
Tubantia agrees with other trachichthyids in the reduction of the neural spine of the
second pre-ural centrum (npu2), the three epurals (ei-3), stegural (st), second uro-
d2
h6
npu2
pul+ul
FIG. 19. Tubantia cataphractus (von der Marck). Caudal skeleton of P. 21984, standard
length 130 mm., Campanian, Baumberg, Westphalia. For explanation of lettering
seep. 102.
neural (d,2], and six hypurals (hi-6). As in Hoplostethus, the fused first ural and pre-
ural centrum (pui + ui] is only as long as the preceding centrum. Tubantia differs
from other Cretaceous trachichthyids and resembles living forms in having the
number of procurrent rays increased to nine spines and two segmented rays above,
six spines and two segmented rays below, these small rays extending forwards in
front of the tips of the neural and haemal spines of the third pre-ural centrum.
Genus GNATHOBERYX Patterson (1967 : 81)
The type and only species, G. stigmosus Patterson (1967 : 82), from the Upper
Santonian of Sahel Alma, Lebanon, is known by two specimens and the caudal
skeleton is preserved only in the holotype, AUB 100402, where it is compressed and
MESOZOIC ACANTHOPTERYGIAN FISHES
75
distorted. So far as can be seen, the caudal skeleton and fin agree with other Creta-
ceous trachichthyids such as Lissoberyx, Hoplopteryx and Acrogaster, with the neural
spine of the second pre-ural centrum reduced, the stegural free, a free second ural
centrum, and the upper principal rays preceded by four spines and one segmented
ray, the foremost articulating with the first epural.
Family HOLOCENTRIDAE Richardson (1846)
The caudal skeleton of the living Myripristis adustus is shown in Fig. 20 : it shows
no significant differences from those of several species of Holocentrus. The neural
and haemal spines of the fourth (puj) and fifth pre-ural centra are expanded but
short. The neural spine of the second pre-ural centrum (npu2) is greatly reduced
2mm
FIG. 20. Myripristis adustus Bleeker. Caudal skeleton of a dried skeleton, 1858 .4.21. 239,
standard length 155 mm., Amboina. hap, hypurapophysis ; for explanation of other
lettering see p. 102.
(and not autogenous as it is in Hoplostethus), with the tip of the first epural lying
above it. The haemal arches of the second and third pre-ural centra (pu2, pu3)
are autogenous and there are three epurals (ei~3) . In contrast to the trachichthyids
the two ural centra, the first pre-ural centrum and the stegural are all fused into a
single structure (pui + UI + U2-}- st) and there are only five hypurals (hi-5)
the uppermost hypural present in trachichthyids having been lost. The second
uroneural (d2) is free and fits proximally into a notch in the stegural rather than
lying below and behind it. In Myripristis the foremost caudal rays articulate with
the first epural above (as in most Cretaceous trachichthyids) and with the haemal
76
THE CAUDAL SKELETON IN
spine of the third pre-ural centrum below. In Holocentrus the foremost rays arti-
culate with the neural and haemal spine of the third pre-ural centrum. In Myrip-
ristis the principal caudal rays are preceded by four spines and one segmented ray
above and below. Rosen (1964, fig. 230) has figured a caudal skeleton of Holo-
centrus ascensionis which differs from all the Recent holocentrid skeletons that I
have seen in having a free second ural centrum. Rosen does not say how large his
specimen was, but if it was an alizarin-stained juvenile this difference can be
explained.
Genus CAPROBERYX Regan (1911 : 8)
In the type species, C. superbus (Dixon) from the Turonian of the English Chalk,
the caudal skeleton is exposed in P. 3979 (Fig. 21). The neural and haemal spines
of the fourth (Puj) and fifth pre-ural centra are normal, not expanded as in living
holocentrids. The haemal arches of the second and third pre-ural centra (pu2,
PUJ) are autogenous, the neural spine of the second pre-ural centrum (npu2) is
reduced and there are three epurals (ei-3), all as in living holocentrids. In contrast
to living holocentrids, the stegural (st) is not fused to the underlying centrum (pui +
ui) and the second ural centrum (u2) is free. The second uroneural (d2) is fused to
the stegural (st) distally, but this is perhaps only a consequence of the very large
size (standard length c. 40 cm) of this specimen. As in living holocentrids there are
only five hypurals, but the distribution of the branched principal fin-rays (one on
e1-3
d2
npu2
10mm
ph
FIG. 21. Caproberyx superbus (Dixon). Caudal skeleton of P. 3979, standard length
c. 400 mm., Chalk, Sussex. For explanation of lettering see p. 102.
MESOZOIC ACANTHOPTERYGIAN FISHES 77
the uppermost hypural, two on the one below and six on the next) shows that the
three upper hypurals are the third and fourth fused (hj + 4), the fifth (/»5) and the
sixth (h6), for in living holocentrids the three upper hypurals, the third, fourth and
fifth, bear two, six and one branched principal rays respectively. Caproberyx
therefore agrees with the trachichthyids in retaining the small sixth hypural, and
the fusion of the third and fourth hypurals shown by P. 3979 is again probably
merely a consequence of the large size of the fish. As in Myripristis and most
Cretaceous trachichthyids, the foremost caudal fin-rays articulate with the first
epural above and the haemal spine of the third pre-ural centrum below, and there
are probably four spines and one segmented ray in front of the principal caudal
rays above and below. Characters of the skull and fins suggest that Caproberyx is the
most primitive holocentrid known, lying near to the common stock of the Holo-
centridae and Trachichthyidae (Patterson 1964 : 359; 1967 : 103). This is con-
firmed by the structure of the caudal skeleton in C. superbus, for in the six hypurals,
the free stegural and second ural centrum, and the unexpanded neural and haemal
spines of the posterior caudal vertebrae, this species resembles the trachichthyids
rather than other holocentrids.
The other species of Caproberyx are C. polydesmus (Arambourg 1954) from the
Lower Cenomanian of Jebel Tselfat, Morocco, and C. pharsus Patterson (1967 : 97)
from the Middle Cenomanian of Hakel, Lebanon. C. pharsus is known only by a
specimen lacking the caudal region. C. polydesmus is known only by the holotype:
Arambourg's figure (1954, pi. 19, fig. i) shows that the neural and haemal spines
of the fourth and fifth pre-ural centra are not expanded, as in C. superbus, and he
described the principal rays as having five or six small rays in front of them in each
lobe.
Genus STICHOCENTRUS Patterson (1967 : 88)
The type and only species is S. liratus Patterson from the Middle Cenomanian
of Hajula, Lebanon. The caudal skeleton is well preserved in AUB 108923 (Fig. 22)
and is partially shown in AUB 108927 and 108929. The neural and haemal spines
of the fifth pre-ural centrum are unmodified but those of the fourth (puj) are
expanded, though not so strongly as in living holocentrids. The autogenous haemal
arches on the second and third pre-ural centra, the reduced neural spine on the
second pre-ural centrum (npu2) and the three epurals (ei~3) are as in Caproberyx
and living holocentrids. The stegural is fused with the underlying centrum (pui +
ui + U2 -\- st) anteriorly. The second uroneural (d2) is free and lies below and
behind the first, not notched into the first as it is in living holocentrids. The second
ural centrum is fused to the compound first pre-ural and ural centrum in AUB
108923 (standard length c. 75 mm.) although the line of fusion is clearly seen, but in
AUB 108929, a much smaller specimen (standard length c. 35 mm.) the centrum
appears free. As in living holocentrids, there are only five hypurals (hi-5), the
small sixth hypural present in Caproberyx having been lost. As in Myripristis,
the foremost caudal rays are inserted on the first epural above and the haemal spine
of the third pre-ural centrum below, and there are four spines and one segmented
ray in front of both the upper and lower principal rays.
THE CAUDAL SKELETON IN
e3
2mm
FIG. 22. Stichocentrus liratus Patterson. Caudal skeleton of AUB 108923, standard
length c. 75 mm., Middle Cenomanian, Hajula, Lebanon. For explanation of lettering
see p. 102. Arrows mark the outermost (unbranched) principal fin-rays.
As in the skull and fins, Stichocentrus is more advanced towards the living
holocentrids than Caproberyx in the expanded neural and haemal spines of the fourth
pre-ural centrum, the five hypurals and the partial fusion of the stegural and second
ural centrum with the preceding centrum.
The remaining Cretaceous holocentrids are Trachichthyoides ornatus Smith Wood-
ward (1902), known only by an isolated head from the English Chalk, and Kansius
sternbergi Hussakof (1929) known by the two syntypes from the Niobrara Formation,
Gove Co., Kansas. Nothing is known of the caudal anatomy of these forms.
Order LAMPRIDIFORMES
Suborder LAMPRIDOIDEI Berg (1940 : 463)
This suborder is used to contain both the Lampridoidei (Lampris only) and the
Veliferoidei (Velifer, etc.] of Berg (Bonde 1966).
? Family VELIFERIDAE Bleeker (1860)
Genus BATHYSOMA Davis (1890 : 424)
The type and only species is B. lutkeni Davis from the Danian stage of southern
Sweden. The caudal skeleton is partially preserved in two specimens in the British
Museum (Natural History). P. 9947 (Fig. 23 A) shows that the upper hypurals are
MESOZOIC ACANTHOPTERYGIAN FISHES
79
fused into a triangular plate and that this plate is fused basally with the second
ural centrum (u2 -\- uh). Above the second pre-ural centrum (pu2) and the fused
first pre-ural and ural centra (pui + ui) there are two elongate bones: the second
of these (e) is certainly an epural but it is impossible to be certain whether the first
is an epural or the neural arch and spine of the second pre-ural centrum. Above
the upper hypural plate there is a third slender bone of uncertain nature. P. 9948
B
pu1+u1
u2+uh
2mm
FIG. 23. Bathysoma lutkeni Davis. Caudal skeleton of A, P. 9947, standard length c.
95 mm.; B, G, P. 9948 (part and counterpart), standard length 78 mm. Both from
Danian, Limhamn, southern Sweden. U2 + uh, second ural centrum fused with one or
more upper hypurals; for explanation of other lettering see p. 102.
(in counterpart, part and counterpart shown in Fig. 236, C) shows the second pre-
ural centrum (pu2) with an autogenous haemal arch and no sign of a neural spine,
the fused first pre-ural and ural centra (pui + ui) bearing the parhypural (ph)
and the first hypural (hi) ,the distal part of the second hypural (h2), and the upper
hypural plate with a fragment of the second ural centrum fused to it (u2 + uh). As
in P. 9947, there is a slender bone lying above the upper hypural plate. The bones
above the first and second pre-ural centra are shattered and displaced. Although
GEOL. 17, 2 7
8o
THE CAUDAL SKELETON IN
these two specimens are far from complete, they show that in Bathysoma the haemal
arch of the second pre-ural centrum was autogenous, the first pre-ural and ural
centra were fused, the parhypural and the first and second hypurals were separate
and autogenous, there was at least one epural, and the upper hypurals were fused
with each other and with the second ural centrum. On the available material it is
difficult to interpret the upper hypural plate and the slender bone above it, which
may be an epural, a stegural or a free hypural. In the upper hypural plate at least
three hypurals can be recognized in transparency under xylene, presumably hypurals
3-5, but the uppermost part of the plate is of a different texture, suggesting that the
stegural may also be fused into the structure, a most unusual condition which can
only be confirmed on more complete material.
e1-3.
st>d2
pti W hp
FIG. 24. Mene maculata (Bloch & Schneider). Caudal skeleton of a dried skeleton,
1866.6.8.59, standard length 95 mm., Taiwan, hp, hypurapophysis ; for explanation
of other lettering see p. 102.
The caudal skeleton of Bathysoma is very different from that of Mene, the only
genus of the Menidae, in which Bathysoma was previously placed (Patterson 1964 :
423). In the living Mene maculata (Fig. 24) the first pre-ural centrum, both the ural
centra, and all but the uppermost hypural are fused into a symmetrical, fan-shaped
plate (pui + ui + U2 + hi-4). The parhypural (ph) has a very large hypura-
pophysis (hp) and lies free below this plate. The uppermost hypural (h^) is also
free, articulating with a hook on the upper edge of the hypural plate. There are
three normal epurals (ei-3) and the neural arch of the second pre-ural centrum
(pu2) is reduced to a very low crest. The stegural (st + d2) is autogenous, articu-
lating with the underlying compound centrum by a large and clearly mobile joint.
The distal part of the shaft of the stegural is grooved longitudinally in M. maculata ;
in the Middle Eocene species M. rhombeus (Volta) and M. oblongus (Agassiz), in
which the caudal skeleton is otherwise identical with that of the living species, there
MESOZOIC ACANTHOPTERYGIAN FISHES 81
is a free second uroneural, and the groove on the shaft of the stegural in M. maculata
clearly marks the line of fusion between the first and second uroneurals. Mene
has seventeen principal caudal rays with fifteen branched, the outermost two or
three rays unsegmented and the inner ones only sparsely segmented, preceded by
five unsegmented but divided (in the median plane) rays above and four below.
The bases of the caudal rays are deeply cleft, covering much of the hypural plate.
The caudal skeleton of Mene can be derived from the basal perciform type (p. 87)
by fusion of the first four hypurals with each other and with the supporting centra,
and the caudal fin has the perciform number of rays. The caudal skeleton of
Bathysoma is of a much more primitive type, differing from that of Beryciformes
mainly in the fusion of the upper hypurals with the second ural centrum. A caudal
skeleton of this type occurs in the Lampridiformes Velifer (Gosline 1961; fig. 3D),
Palaeocentrotus (Kiihne 1941, fig. 2 ; Bonde 1966) and Lampris, and the known skele-
tal features of Bathysoma (Patterson 1964, fig. 90) agree as well with Palaeocentrotus
and Velifer (Regan 1907; Smith 1951) as they do with Mene. The holotype of
Bathysoma lutkeni, in Copenhagen, shows that the supraoccipital crest is attached
to the skull roof only at the posterior end, with a gap between it and the frontal
crest (Bonde, personal commn) : an exactly similar supraoccipital crest occurs in
Bonde's (1966) ? veliferid from the Lower Eocene [Mo-clay], while in Palaeocentrotus
there is a large foramen between the supraoccipital and frontal crests (Kiihne 1941,
fig. 3). Provisionally Bathysoma may be placed in the Veliferidae but, like Bonde's
Eocene form, it may well prove to be closer to Palaeocentrotus.
Order SALMONIFORMES (Greenwood et al. 1966)
Suborder MYCTOPHOIDEI
The most generalized of living myctophoids is Aulopus (Aulopodidae, Regan
191 la : 121 ). The caudal skeleton of Aulopus is briefly discussed by Gosline
(1961 : 10), who notes that this genus is one of the few living teleosts retaining large
caudal scutes in front of the caudal lobes. The caudal skeletons of more advanced
myctophoids have been figured by HoUister (Synodus, Trachinocephalus , 1937^,
figs 1-14), Gosline (C hlorophthalmus , 1961, fig. 26), Rosen (Myctophum, 1964, fig. 236)
Greenwood et al. (Neoscopelus, 1966, fig. 36) and Weitzman (Parasudis and Saurida
1967, figs 17, 18). In Aulopus the caudal skeleton is almost identical with that of the
Cretaceous Ctenothrissa (Figs 4, 5), with a free second ural centrum, a stegural and a
second uroneural, three epurals and six hypurals, of which the first and third are
the largest. The only differences from Ctenothrissa are that the haemal arch of the
third pre-ural centrum and the neural arch of the second pre-ural centrum are auto-
genous, the neural spine of the second pre-ural centrum is less expanded, more
spine-like, and just over half as long as its predecessor, and the second ural centrum
has a long posterior process. In all these characters, Aulopus appears to be more
primitive than Ctenothrissa. In other living myctophoid families conditions are
much as in Aulopus, but the caudal scutes are lost, the neural spine of the second
pre-ural centrum ceases to be autogenous and becomes shorter and expanded, the
first epural tending to move forwards above it, there is often fusion within the upper
82
THE CAUDAL SKELETON IN
and lower hypurals, the sixth hypural and one epural may be lost (Synodontidae),
and the second ural centrum and stegural may fuse with the compound first ural
and pre-ural centrum. These trends are very like those seen within the Berycoidei
(p. 68; cf. figs. 236, D in Rosen 1964).
In the Upper Cretaceous myctophoids were abundant, the best known genera
being Sardinioides, Acrognathus, Cassandra ( = Leptosomus) and Nematonotus.
Nematonotus appears to be the most primitive of these and will serve as an example.
Figure 25 shows specimens of Nematonotus bottae (Pictet & Humbert), from the
Cenomanian of Hakel, Lebanon, and N. longispinus (Davis), from the Cenomanian
e1-3
d2-
e1-3.
h5
pu1*u1
pu1*u1
ph
ud
2mm
FIG. 25. The caudal skeleton in A, Nematonotus bottae Pictet & Humbert, 49563, stand-
ard length c. 90 mm., Middle Cenomanian, Hakel, Lebanon; B, Nematonotus longispinus
(Davis), P. 13882, standard length 114 mm., Middle Cenomanian, Hajula, Lebanon.
In G are the bases of the upper caudal rays in P. 48825, N. longispinus, standard length
97 mm., to show the urodermal, ud. For explanation of other lettering see p. 102.
In B the second uroneural is missing, in A and C arrows mark the outermost (unbranched)
principal fin-rays.
MESOZOIC ACANTHOPTERYGIAN FISHES 83
of Hajula, Lebanon. There are no significant differences between these two species
in the caudal region. In the specimen of N. bottae illustrated, the haemal arches
of the third and fourth pre-ural centra are fused and the haemal spine of the third
pre-ural centrum is partially doubled. This is an individual abnormality. As in
Aulopus there is a large caudal scute (/. s) above and below the caudal skeleton and
the neural spine of the second pre-ural centrum (npu2) is autogenous, spine-like and
just over half as long as its predecessor. The autogenous haemal spines on the
second and third pre-ural centra, three epurals (ei-3), six hypurals (hi-6), long second
ural centrum (u2, Fig. 256) and second uroneural (d2) are as in Aulopus. The
stegural (st) is forked proximally, with a process extending forwards on to the second
pre-ural centrum. This process, absent in Aulopus and Ctenothrissiformes, is a
primitive feature present in Flops and some clupavids (Patterson 19670, fig. n)
which indicates the double origin of the first uroneural (Regan 1910 : 355 ; Patterson
1968 : 226). In Nematonotus there is a single urodermal (Fig. 256), a structure absent
from living myctophoids and Ctenothrissiformes but also present in the Cenomanian
Sardinioides attenuatus. In Nematonotus the foremost procurrent caudal rays
articulate with the neural and haemal spine of the third pre-ural centrum and the
nineteen principal rays are preceded by four unsegmented and four segmented rays
above, four unsegmented and two segmented rays below. In myctophoids the
procurrent caudal rays are normally longitudinally divided, not spinous, but pro-
current caudal spines appear in advanced members of the family Myctophidae
(Fraser-Brunner 1949 : 1033).
III. DISCUSSION
(a) The relationships of Ctenothrissiformes, Myctophoidei and Beryciformes.
Monod (1967 : 118) has remarked that the structure of the caudal skeleton is
rarely of value in discriminating between taxa at the generic level and below, but
that it becomes increasingly valuable at the familial, subordinal and ordinal level.
This observation is fully borne out by the forms described here, the only exception
being the occurrence of two types of caudal skeleton in the ctenothrissiform family
Aulolepididae, Pateroperca having the neural spine of the second pre-ural centrum
fully developed, Aulolepis having it short and expanded, as it is in Ctenothrissidae.
This suggests that Aulolepis and Pateroperca belong to different families, but
Pateroperca is as yet so poorly known that no conclusion can be reached on this
point until more specimens are discovered.
Among the Ctenothrissiformes, Myctophoidei and Beryciformes caudal anatomy
is very similar. The basal type of caudal skeleton in these groups has the following
features: the first pre-ural and ural centra fused, a free second ural centrum, three
epurals, a stegural, a second uroneural and six hypurals (two lower and four upper).
Caudal scutes above and below the caudal skeleton are present in Aulopus, most
Cretaceous myctophoids (Nematonotus, Acrognathus, Sardinioides) and Ctenothriss-
iformes (Aulolepis, Ctenothrissa) : they are absent in Beryciformes and higher groups.
A single urodermal is present in Nematonotus and Sardinioides attenuatus among
Cretaceous myctophoids. There is no urodermal in Ctenothrissiformes or Bery-
ciformes. The main variations encountered in the caudal skeletons of myctophoids,
GEOL. 17, 2 7§
84 THE CAUDAL SKELETON IN
ctenothrissiforms and beryciforms involve reductions in the number of caudal
elements by fusion or suppression and the condition of the neural spine of the second
pre-ural centrum, which may be fully developed (Pateroperca, Polymixioidei, Pycno-
steroides, Dinopteryx), spine-like and about half as long as its predecessor (Nematono-
tus, Aulopus, Aipichthys, Pharmacichthys) , short and expanded (Ctenothrissa,
Aulolepis, many myctophoids) or reduced to a low crest (Berycoidei) .
The basal teleostean caudal skeleton, seen in such genera as Leptolepis, Allothris-
sops, Thrissops, Ichthyodectes, Hiodon, Flops, Salmo, etc., contains two free ural
centra, three epurals, two lower hypurals and five upper hypurals (there may be six
or seven upper hypurals in Leptolepis; Patterson 1968 : 220; there are only four in
salmonids; Norden 1961 : 738), up to seven uroneurals (Patterson 1968) which extend
forwards to the fourth pre-ural centrum in Thrissops and Ichthyodectes, to the third
in Leptolepis and Allothrissops and to the second pre-ural centrum in Hiodon, Elops,
Salmo, etc., and one or two urodermals (Patterson 1968 : 230). The condition of the
neural arches and spines in the caudal region is often complicated by doubling of the
segmental structures, usually as individual variations, but in Leptolepis, Allothrissops,
Flops and some salmonids there is normally a neural arch and spine on the first pre-ural
centrum, and in Leptolepis, Hiodon, osteoglossoids (Greenwood 1967) and Alepo-
cephalus (Patterson 1968, fig. 12) there may be a more or less well developed arch and
spine on the first ural centrum. It is usually assumed (e.g. Gosline 1961 : 14; Patterson
19670 : 104) that the second pre-ural centrum primitively bears a complete neural
spine, supporting epaxial fin-rays. But among the forms described here the most
primitive (Aulopus, Nematonotus) have the neural spine of the second pre-ural
centrum about half as long as that of the third, and this is also true of Elops (Nybelin
1963, figs, i, 4), which in other respects seems to be the most primitive living teleost.
This suggests that the short second pre-ural neural spine may be primitive for some
teleost groups. There are three possible conditions of the second pre-ural neural
spine. It may be fully developed, as in Allothrissops (Patterson 19670, fig. 6),
Ichthyodectes (Cavender 1966, fig. i), Tarpon (Nybelin 1963, fig. 7), and many other
primitive telosts, normally supporting epaxial fin-rays but in Tarpon ending just
in front of the foremost epaxial fin-ray; it may be about half as long as its prede-
cessor, as in Elops, " Clupavus " (Patterson 19670, fig. n), Nematonotus and Aulopus;
or it may be represented only by a low crest, as in Berycoidei and generalized percoids
(p. 87). Intermediates between the second and third of these conditions occur in
Aulolepis, Ctenothrissa and many myctophoids. Intermediates between the first
two conditions seem to occur only in primitive protacanthopterygian groups
(salmonids, as in the specimen of Cristivomer illustrated by Vladykov 1954, fig. 2;
characinids, as in the specimen of Brycon illustrated by Weitzman 1962, fig. 15)
in which the pre-ural neural spines are very variable and both conditions may occur
in a single species. The third condition, the spine reduced to a low crest, is
undoubtedly advanced and may be left out of consideration here. In the ancestors
of the teleosts, the pholidophorids, the neural spines of the last three pre-ural verte-
brae decrease in size progressively so that all three end on approximately the same
oblique plane : the first pre-ural neural spine is very short, the second is both shorter
and more slender than the third (Patterson 1968, figs. 1-4). In pholidophorids
MESOZOIC ACANTHOPTERYGIAN FISHES 85
these neural spines do not reach the dorsal edge of the trunk and do not support
fulcra or fin-rays. This condition of the pre-ural neural spines seems to be primitive
for the teleosts as a whole and it persists in the Lower Jurassic Leptolepis cory-
phaenoides and L. normandica (Nybelin 1963, figs. 9, 10) and the Upper Jurassic
L. dubia (Nybelin 1963, fig. 8; Patterson 1968, fig. 10). In the Upper Jurassic
two of the three modern types of second pre-ural neural spine were already in exis-
tence. In Allothrissops and Thrissops (Nybelin 1963, figs, u, 12; Patterson 19670,
fig. 6) both the second and third preural neural spines have elongated so that they
reach the dorsal edge of the trunk, ending just in front of the foremost procurrent
fin-rays: this is essentially the condition in living Tarpon. In the Upper Jurassic
E lops-like fish illustrated by Nybelin (1963, fig. 6) the second pre-ural neural spine
remains short but the third and fourth pre-ural neural spines are elongated, reaching
the dorsal edge of the trunk and supporting the foremost procurrent fin-rays. This
is essentially the condition in living Flops. There is no a priori reason to regard
either of these two conditions as more primitive, both are a response to a new need,
the necessity to support the epaxial procurrent rays as they extend forwards to
increase the dorso-ventral symmetry of the tail. However, we know that the Elops
type, with a short second pre-ural neural spine, has persisted unchanged in elopids
since the Upper Jurassic and that this type of second pre-ural neural spine is primi-
tive for the teleosts as a whole, and there is no reason for regarding the short neural
spine of Elops and its Jurassic relative as a secondary regression from a long neural
spine of Tarpon type. I conclude, therefore, that when one finds a second pre-ural
neural spine resembling that of Elops in a generalized teleost one should regard it
as a primitive feature unless there is good evidence to the contrary. In support
of this interpretation is the occurrence of a second pre-ural neural spine of this type
only in teleosts in which the caudal skeleton retains such primitive features as a
free second ural centrum and nineteen principal caudal rays. A generalized teleost
having an elongate second pre-ural neural spine is to be regarded as having developed
this from a short spine of leptolepid or elopid type : this development can apparently
take place spontaneously (see Aipichthys, Fig. n).
From the basal type of teleostean caudal skeleton, the most primitive members of
the myctophoid-ctenothrissiform-beryciform assemblage (such as Nematonotus}
differ in the loss of one hypural, the seventh, whether by suppression or by fusion
with the sixth is as yet unknown, have reduced the number of uroneurals to two,
principally by loss of the small posterior uroneurals (ural neural arches 6-8), since
the forked first uroneural of Nematonotus (Fig. 25) is clearly homologous with that
of Elops, representing the second and third ural neural arches, while the second
uroneural is probably homologous with the second uroneural of Elops, representing
the fourth and fifth ural neural arches (Patterson 1968 : 226), the first pre-ural and
ural centra have fused, and the first uroneural has fused with the first ural and pre-
ural neural arches to produce a stegural, a development which took place very early
in the protacanthopterygian lineage (salmonids, " Clupavus ", etc.). These changes
raise the question of the origin of the myctophoids and ctenothrissiforms. Green-
wood et al. (1966 : 371) wrote of the ctenothrissiforms " we link them with some
early group of myctophoid-like salmoniform fishes in which the supramaxillae were
86 THE CAUDAL SKELETON IN
not reduced, the premaxilla had not excluded the maxilla from the gape, and in
which the adipose fin had disappeared. " By extension, the ancestor of the mycto-
phoids would be such a fish with an adipose fin. On the other hand, Gosline (1961 :
35 ; also Gosline, Marshall & Mead 1966 : 5) points out that the large caudal scutes
of Aulopus make it impossible to derive the myctophoids from any living teleost
except the elopoids. Weitzman (1967 : 532) discusses this point and notes that
caudal scutes are present in Argentina, but the structures he describes in the salmonoid
Plecoglossus and the galaxioid Retropinna do not seem to resemble caudal scutes.
The absence in all known salmonoids of large caudal scutes and of a forked first
uroneural of the type found in Elops and Nematonotus, together with the absence of
recognizable salmonoids from pre-Tertiary rocks, make it difficult to envisage any
direct relationship between salmonoids and myctophoids (Greenwood et al. 1966, fig. i) .
A short second pre-ural neural spine resembling those of Aulopus and Nematonotus
occurs in some salmonoids (Coregonus, which also has a urodermal), but the last few
neural arches and spines and the epurals are apparently very variable, and no clear
pattern emerges from Norden's (1961 : 738) analysis. In Argentina caudal scutes are
present (though they are reduced) and the first pre-ural and ural centra are fused
(Gosline 1960, fig. 10), as they must have been in the common ancestor of Myctophoidei
and Ctenothrissiformes, but the neural spine of the second pre-ural centrum is elon-
gate and supports procurrent fin-rays : in this character Argentina is advanced over the
basal myctophoids and ctenothrissiforms. In the Clupavidae, a family with a fossil
record extending back to the Upper Jurassic, large caudal scutes are usually present
in the tail, the skull seems primitive enough to have given rise to both myctophoids
and Ctenothrissiformes, and the caudal skeleton may be strikingly like that of
Nematonotus (Patterson 19670, fig. n). It is unlikely that the known clupavids
were ancestral to the myctophoids and ctenothrissiforms because of their reduced
dentition and clupeid-like jaws, with a high coronoid process on the dentary, but
they suggest a possible source for this type of caudal skeleton.
Taking the caudal skeleton of Nematonotus as the primitive condition for the mycto-
phoids and ctenothrissiforms, the myctophoids are characterized by shortening
and broadening the neural spine of the second pre-ural centrum, and in more
advanced forms by fusion of the hypurals with each other and with the supporting
centra. Within the ctenothrissiforms two distinct types of caudal skeleton occur:
in Ctenothrissa and Aulolepis the neural spine of the second pre-ural centrum is
expanded, as in myctophoids, tending towards the condition in Berycoidei and
Percoidei; in Pateroperca the second pre-ural neural spine is fully developed, as it
is in Polymixioidei. In Aulolepis and some species of Ctenothrissa there are pro-
current spines in front of the caudal fin : this is specific evidence of evolution towards
the Berycoidei and like the reduction of the second pre-ural neural spine in these
fishes it distinguishes them from the Polymixioidei and Dinopterygoidei, in which
procurrent caudal spines occur only in advanced forms.
Within the Beryciformes, the structure of the caudal skeleton gives some support
to the division of the order into three suborders. In Polymixioidei there are always
six hypurals and the neural spine of the second pre-ural centrum is fully developed.
In Berycoidei the second pre-ural spine is reduced to a low crest (except in one
MESOZOIC ACANTHOPTERYGIAN FISHES 87
individual of Monocentris, Fig. 14), the procurrent rays are spinous, and there are
trends towards loss of the sixth hypural and fusion of the stegural and second ural
centrum with the preceding centrum. In the Dinopterygoidei, already known to be
a heterogeneous group, the neural spine of the second pre-ural centrum retains the
primitive short condition (Aipichthys, Pharmacichthys) or is fully developed (Dinop-
teryx, Pycnosteroides) and there are trends towards loss of the sixth hypural. The
polymixioid caudal skeleton (also found in Dinopteryx and Pycnosteroides) resembles
that of the ctenothrissiform Pateroperca, differing only in having no caudal scutes
and in having one less principal ray. The caudal skeleton of Berycoidei resembles
those of Aulolepis and Ctenothrissa, differing only in having lost the caudal scutes
and further reduced the second pre-ural neural spine. Aipichthys and Pharma-
cichthys seem to have the most primitive caudal skeletons known in Beryciformes,
with nineteen principal rays, the primitive short second pre-ural spine, and, at least
in some specimens of Aipichthys, six autogenous and separate hypurals and a free
second ural centrum. Although both Aipichthys and Pharmacichthys are too
specialized in other characters to have given rise to Beryciformes, their caudal skele-
ton could give rise to both the polymixioid condition (by elongation of the second
pre-ural neural spine, which occurs spontaneously in some individuals of Aipichthys,
Fig. nC) and the berycoid condition (by shortening of the second pre-ural neural
spine).
(b) The origin ofPerciformes.
Gosline (i96ia) discussed the caudal skeleton of Perciformes and found that the
most generalized type contains fifteen branched principal rays, no neural spine on the
second pre-ural centrum, three epurals, two free uroneurals, no free ural centra,
five autogenous hypurals, and the haemal arches of the second and third pre-ural
centra autogenous (see also Monod 1967, fig. 3). Gosline mentioned that this
type of caudal skeleton occurs in Kuhlia (Kuhliidae), Chaetodon (Chaetodontidae),
Polydactylus (Polynemoidei) and juvenile Sphyraena (Sphyraenoidei) . Monod (1967)
refers to this type of caudal skeleton as " sciaeno-sparidien banal " and states that
it occurs in many Perciformes, mentioning Sciaena (Sciaenidae), Pagrus, Sparus
(Sparidae) and Gaterin (Pomadasyidae) . I find that this generalized type of caudal
skeleton also occurs in Centropomus (Centropomidae) , Lateolabrax, Polyprion,
Dicentrarchus, M or one, Acanthistius (Percichthyidae, sensu Gosline 1966), Branchio-
stegus (Branchiostegidae), Pomatomus (Pomatomidae), Brama (Bramidae), Arripis
(Arripidae), Lutjanus (Lutjanidae), Nemipterus, Scolopsis (Nemipteridae), Lobotes
(Lobotidae), Xenocys, Xenistius, Xenichthys (Pomadasyidae), Lethrinus, Sphaerodon
(Lethrinidae), Monodactylus (Monodactylidae) , Kyphosus, Medialuna (Kyphosidae) ,
Ephippus, Drepane, Platax (Ephippidae), Chelmo, Heniochus, Pomacanthus (Chaeton-
ontidae), Histiopterus (Pentacerotidae), Cirrhitus (Cirrhitidae) and Schedophilus
(Stromateoidei). Many other groups, among them the Serranidae (sensu Gosline
1966), Cichlidae, Percidae, Acanthuroidei, etc. differ from this basal type only in
the loss or incorporation in the stegural of the second uroneural. The occurrence
of an apparently identical type of caudal skeleton in such a wide range of perciform
groups, including forms with lunate, forked, emarginate and rounded caudal fins
88 THE CAUDAL SKELETON IN
in habitats ranging from pelagic to lacustrine, suggests that caudal anatomy
is unlikely to contribute much to the unravelling of lineages among generalized
Perciformes.
A caudal skeleton very similar to the basal perciform type occurs in some advanced
Berycoidei (living Holocentridae and Diretmidae differ only in having the stegural
fused with the underlying centrum (Fig. 20), Berycidae have the stegural fused with
the centrum and also retain the sixth hypural (Regan 1911, fig. i)) but here the
hypurals support nineteen principal rays. In Myctophoidei a slightly different
sequence of fusion is followed in which the second ural centrum partially retains its
individuality. So far as I know, the basal perciform caudal skeleton is not precisely
duplicated elsewhere.
The differences between the caudal skeleton and fin of basal Perciformes and those
of generalized Beryciformes (Polymixia, Aipichthys, Pycnosteroides, Monocentris) are:
(i) Reduction of the neural spine of the second pre-ural centrum. This has
already taken place in all Berycoidei.
(ii) Fusion of the second ural centrum with the preceding two centra. This has
taken place in most living Berycoidei and occurs in some individuals of Aipichthys.
(iii) Loss of the sixth hypural (the development of the caudal skeleton in Mugil
and Sphyraena (Hollister 1937) suggests that Perciformes have lost the sixth hypural,
not incorporated it in the fifth). This has already occurred in some individuals of
Aipichthys and Pycnosteroides, and takes place during the evolution of the Berycoidei
(Holocentridae, Diretmus] .
(iv) Reduction of the number of principal caudal rays from nineteen (Berycoidei,
Aipichthys and Pharmacichthys] or eighteen (Polymixioidei, Dinopteryx and Pycnos-
teroides) to seventeen.
(v) In all living Beryciformes, the foremost procurrent rays in each caudal lobe
are true spines, but in Perciformes they are usually (? always) unsegmented lepido-
trichia, with the right and left halves separate, and this is true of the earliest Perci-
formes (Prolates). In this character Perciformes are more primitive than living
Beryciformes. Among Cretaceous Beryciformes, all Berycoidei, like their living
relatives, have spines in front of the caudal fin; in Polymixioidei procurrent spines
occur only in Homonotichthys and Pycnosterinx dubius, already known to be evolving
towards the living Polymixia (Patterson 1964 : 301, 380), and in Dinopterygoidei
they occur only in Dinopteryx. It is striking to find that spinous procurrent caudal
rays occur only in those Cretaceous genera (except Dinopteryx} already known to be
closely related to riving Beryciformes.
The first of these five differences, the condition of the neural spine of the second
pre-ural centrum, is the most interesting. As discussed above (p. 84) the primitive
condition of this structure in teleosts seems to be as in Flops, Aulopus and Nematono-
tus, where the spine is slender and about half as long as its predecessor. This
type of spine may elongate so that it supports procurrent fin-rays, as in Polymixioidei
and many primitive telosts, or it may become reduced to the percoid condition (Fig.
28) . But apart from these two simple alternatives there are other possibilities which
complicate the issue. First, the fully developed spine might become detached as an
epural, producing the percoid condition direct. Secondly, from a low neural crest
MESOZOIC ACANTHOPTERYGIAN FISHES 89
of percoid type an apparent full neural spine might develop secondarily by fusion
with the first epural. Thirdly, an apparent neural spine on the second pre-ural
centrum might be produced by fusion between the second and third pre-ural centra.
The last of these possibilities can normally be recognized by the partial or complete
doubling of the neural or haemal spine on the compound centrum, as in Pleuronectes
(Harrington 1937, fig. i) and the specimen of Saurida illustrated by Weitzman
(1967, fig. 18). The abnormal specimen of Monocentris illustrated in Fig. 14 is
evidently a special case of this type of fusion, where the neural and haemal spines
of the third pre-ural centrum are normally double and the posterior half of the
neural spine has become attached to the succeeding centrum. This type of fusion
does not seem of general significance in the present discussion.
Fusion of an epural with the neural crest of the second pre-ural centrum to produce
a secondary neural spine (Fig. 28E) is a common occurrence in acanthopterygians :
this process appears to account for the complete neural spine on the second pre-ural
centrum in such groups as the Nandidae (but not Pristolepis ; Gosline 1968, fig. 2b)
among Percoidei, the Channiformes, Anabantoidei, Luciocephalus , some scombroids,
pleuronectoid and soleoid pleuronectiforms, tetraodontiforms, etc. (Monod 1967;
Liem 1963, 1967; Gosline 1968). In Psettodes, the most primitive living pleuronecti-
form, the caudal skeleton (Monod 1967, fig. 13) is of basal perciform type, with five
autogenous hypurals and two uroneurals, but there is only one free epural and there
appears to be a neural spine on the second pre-ural centrum. Monod identifies this
spine as the first epural, for the element is partially or completely autogenous and
the suture at the base lies not between the arch and the centrum but between the
arch and the spine. Psettodes demonstrates clearly that the neural spine of the
second pre-ural centrum in Pleuronectoidei and Soleoidei is an epural which has
secondarily regained contact with and fused with a neural arch. That this has also
occurred in scombroids such as Neothunnus can be seen by comparing figs. 15 and
16 of Monod (1967) . A further peculiarity of the caudal skeleton of pleuronectoids
and soleoids is that the parhypural tapers proximally and fails to make contact with
the centrum (Monod 1967 : 117). The effect of this is to give dorso- ventral sym-
metry to the caudal skeleton, the free parhypural opposing the single epural just as
the neural and haemal spines of the second pre-ural centrum oppose one another.
A free parhypural, tapering proximally, also occurs in acanthopterygians such as
the Channiformes (Monod 1967 : 117; Gosline 1968, fig. 20), most Anabantoidei
(Liem 1963 : 32), Luciocephalus (Liem 1967 : 114) and balistoids (Whitehouse 1910,
pi. 50, fig. 33; Monod 1967 : 117), all forms with a neural spine on the second pre-
ural centrum. Since none of these fishes has more than two epurals, all these groups
appear to be cases of secondary fusion between the first epural and the second pre-ural
neural arch in order to increase the dorso-ventral symmetry of the caudal skeleton.
A complete neural spine on the second pre-ural centrum also occurs occasionally in
basal percoids: Fig. 26 shows such a structure in a large specimen of Siniperca
(Percichthyidae). In this individual there is a perfectly formed neural arch and
spine (npu2] fully fused to the second pre-ural centrum, and there are only two
epurals (ei, 2] compared with the three of normal Siniperca, most percichthyids and
serranids. This specimen is best regarded as an abnormality foreshadowing the
go
THE CAUDAL SKELETON IN
fusion of the first epural with the second pre-ural neural arch in nandids, pleuro-
nectiforms, etc.
It is thus well established that in many perciform groups and perciform derivatives
the first epural can fuse with the second pre-ural neural arch. This raises the
question of the homology of the perciform first epural: does the frequent fusion of
this bone with the second pre-ural centrum indicate that these two structures were
h5.
FIG. 26. Siniperca chuatsi (Basilewsky). The caudal skeleton of a dried skeleton showing
a complete neural spine on the second pre-ural centrum, 1888.3.23.3, standard length
340 mm., Kiu Kiang, China. For explanation of lettering see p. 102.
originally part of the same segment? If so, the perciform first epural may have
appeared by detachment of a fully developed neural spine in a caudal skeleton of
polymixiid type. Rosen (1964 : 244) suggested that this took place in the evolution
of the exocoetoids : that the ancestral exocoetoid had only two epurals and a fully
developed neural spine on the second pre-ural centrum which became detached as
the foremost of the three epurals in such a fish as Dermogenys. If this neural spine
became detached in a fish which still retained the original three epurals one would
expect there to be four epurals. Such a condition occurs very occasionally in
MESOZOIC ACANTHOPTERYGIAN FISHES
.d2
h5
•puUuUu2
5 mm
FIG. 27. Pentaceropsis recurvirostris (Richardson). Caudal skeleton of a dried skeleton
showing four epurals, 1869.2.24.24, standard length 405 mm., Tasmania. For explana-
tion of lettering see p. 102.
the generalized percoid caudal skeleton, as in the individual of Pentaceropsis
(Pentacerotidae) shown in Fig. 27.
The epurals of teleosts are the remnants of a more numerous set of bones in primi-
tive actinopterygians which are serial homologues of the supraneurals above the
anterior vertebrae, and like these bones they were primitively metameric (Patterson
1968 : 221), but the metameric arrangement of the epurals has been lost in living
chondrosteans. In the most primitive teleosts, such as the Jurassic Leptolepis and
Allothrissops (Patterson 19670:, figs. 3, 6), the three epurals still show an apparent
metamery, suggesting that they represent the neural spines of the first pre-ural
neural arch and two ural neural arches : this seems to be supported by conditions in
osteoglossoid fishes, where there are often fully-developed neural spines on both
the first pre-ural and the first ural centra, and where there is never more than one
epural (Greenwood 1967). But in higher teleosts, as in living chondrosteans, a
metameric arrangement of the epurals is no longer recognizable. At the percoid
level, for example, one can find fishes with all the epurals behind the neural crest on
the second pre-ural centrum (Gosline ig6ia, fig. i, Kuhlia; 1968, fig.^c.Bathymaster),
or with one epural above the crest (Gosline ig6ia, fig. 2, Parupeneus; Hollister 1937,
92 THE CAUDAL SKELETON IN
figs. 12-14, Sphyraena) or with two above it (Hollister 1937, fig. 8, Mugil). Further,
in groups such as the Berycoidei and Percoidei there is good evidence that in primi-
tive forms the first epural lies behind the neural crest of the second pre-ural centrum,
moving forwards above it in more advanced forms (in berycoids cf. Figs. 17, 21 with
Figs. 13, 20). Since there is no evidence that strict metamery of the epurals is
maintained in acanthopterygians there is no reason to believe that the perciform
first epural is the detached neural spine of the second pre-ural centrum. The
individual of Pentaceropsis shown in Fig. 27 is best interpreted merely as exhibiting
a supernumary epural: such a condition is already known to occur in the salmonid
Oncorhynchus, where Vladykov found four epurals in three out of 1,020 specimens
(1962, tableS).
The conclusions drawn from this discussion of the second pre-ural neural arch
and spine are that the low neural crest in basal percoids is to be regarded as having
evolved by reduction of the primitive short neural spine, and that the ancestors of
the perciforms are to be found among fishes having a low crest or short neural spine
on the second pre-ural centrum, not among those with a full neural spine. Where
there is a full neural spine on the second pre-ural centrum in Perciformes and perci-
form derivatives, it is to be regarded as having arisen by fusion of an epural with a
low neural crest. These conclusions are illustrated in Fig. 28.
With this background on the perciform caudal skeleton, we can now consider the
evidence of caudal structure in the various beryciform-perciform lineages that have
been suggested (Patterson 1964). These were, in decreasing order of confidence,
Polymixiidae (Omosoma — Berycopsis lineage) — — > Scorpididae,
Monodactylidae and
Kyphosidae
Aipichthyidae — > Carangidae
Pharmacichthyidae — > Acanthuroidei
Sphenocephalidae — > basal Percoidei
(Serranidae, etc.)
Pycnosteroididae — > Chaetodontidae
Dinopterygidae — > Centrarchidae
Detailed study of the caudal skeleton cannot be said to give support to these
lineages. Of the various perciform groups mentioned, the scorpidids, monodacty-
lids, kyphosids, basal percoids (Centropomidae, Percichthyidae), Chaetodontidae
and Centrarchidae have the generalized percoid type of caudal skeleton, differing
from those of Beryciformes in the characters listed on p. 88. The acanthuroids
are more advanced only in having lost the second uroneural, while the carangids
have enlarged the first epural and show fusion between the first and second hypural
and between the third and fourth hypurals. Among the beryciform groups, the
polymixiids, sphenocephalids, pycnosteroidids and dinopterygids all have a com-
plete neural spine on the second pre-ural centrum : for reasons given in the discussion
above, it is unlikely that a percoid caudal skeleton can be derived directly from this
condition. In Sphenocephalus there are only two epurals, suggesting the possibility
MESOZOIC ACANTHOPTERYGIAN FISHES
93
of arriving at a percoid arrangement of three epurals and a low neural crest on the
second pre-ural centrum by detachment of the neural spine on this centrum, but
further study of Sphenocephalus has yielded strong evidence (to be discussed in
a forthcoming paper by the author and D. E. Rosen) that far from being an ancestral
percoid it is related to the percopsiforms. In the aipichthyids and pharmacichthyids
FIG. 28. Diagrams to show changes in the second pre-ural neural spine (black) in the
evolution of the myctophoids, ctenothrissiforms and acanthopterygians. A, the primi-
tive short neural spine, as in Nematonotus, Aulopus, Aipichthys and Pharmacichthys;
B, elongation of the neural spine to support procurrent fin-rays, as in Pateroperca, Poly-
mixiidae, Dinopteryx and Pycnosteroides ; C, expansion of the neural spine into a plate, as
in most myctophoids, Ctenothrissa, A ulolepis and some primitive Berycoidei ; D, reduction
to a low crest, with the first epural (stippled) moving forwards above it, as in most
Berycoidei and Perciformes (a berycoid is illustrated) ; E, production of a secondary neural
spine by fusion of the first epural in fishes with no free second ural centrum and five
hypurals, as in Zeiformes (except Caproidae), Nandidae, Channiformes, Anabantoidei,
Pleuronectiformes, etc.
94 THE CAUDAL SKELETON IN
the neural spine of the second pre-ural centrum is still short, as in the most
primitive teleosts, and could give rise to the percoid condition by reduction. Further,
in Aipichthys (Fig. n) there is a tendency to reduce the number of hypurals to five
and also to fuse the second ural centrum into the preceding centrum, as in percoids.
There seems to be nothing in the caudal skeleton of Aipichthys to oppose the postu-
lated link with the carangids. In Pharmacichthys the caudal skeleton is like that
of Aipichthys, so far as it is known, and the fin-rays are deeply cleft basally, as in
carangids: this character reduces the possibility of a relationship between Phar-
macichthys and the acanthuroids and balistoids, in which the caudal fin-rays are
unmodified.
Except for the Aipichthys-carangid lineage, the evidence of the caudal skeleton
indicates that Gosline's (19660) criticism of the polyphyletic scheme of perciform
origins which I proposed are well founded. But the evidence on which Gosline
based these criticisms, the structure of the supraoccipital crest, does not in fact
oppose the various beryciform-perciform lineages. Gosline (19660 : 412) contrasted
the type of supraoccipital crest seen in the polymixiids Homonotichthys and Poly-
mixia, and in the carangids, priacanthids, etc., which extends forwards between the
frontals, is knife-edged and buried in musculature, with the type of crest seen in
Antigonia, acanthuroids, chaetodontids, etc., which is short, high and thickened
anteriorly, extending " up and back over the nape as a sort of protective shell ".
Gosline finds that these two types of supraoccipital crest are " structurally and
functionally ... far apart ", and writes " nor does it appear that one could be
developed from the other except by going all the way back through some inter-
mediate form with a relatively small, unspecialized occipital crest ". Gosline draws
the conclusion that Aipichthys and Sphenocephalus , both with a short crest, thickened
anteriorly, could not have given rise respectively to the carangids and serranids,
which have a long, knife-edged crest. But the evolution of the Berycoidei shows
(as clearly as such processes can be shown by the fossil record) that fishes with a
short " Aipichthys-type " crest can give rise to fishes with a long " carangid-type "
crest. The most primitive Berycoidei have an " Aipichthys-type, " crest, moderately
high and thickened anteriorly : this is true of both the trachichthyid lineage (Lisso-
beryx, Patterson 1967, fig. 2) and the holocentrid lineage (Caproberyx, Patterson
1964, fig. 67; 1967, fig. 10, Stichocentrus, Patterson 1967, fig. 8). From this basal
type there are in berycoids two divergent trends in the evolution of the supra-
occipital crest. In Holocentridae the frontals grow backwards, partially covering
the parietals, eliminating the supratemporal fossa, and producing a low supra-
occipital crest which secondarily comes to resemble the small supraoccipital crest
of primitive teleosts. In Trachichthyidae (Hoplopteryx, Hoplostethus) the supra-
temporal fossa extends forwards and the supraoccipital crest becomes elongated,
thickened centrally and knife-edged. This trend continues further in the Berycidae,
and Beryx has a long knife-edged crest, continued forwards by the frontals above the
orbit, which resembles those of Polymixia and the carangids. If a carangid-
type " supraoccipital crest developed from an " Aipichthys-type. " within the
Berycoidei, there is no reason why these changes could not have occurred in other
lineages.
MESOZOIC ACANTHOPTERYGIAN FISHES 95
It should also be mentioned here that Gosline (19660 : 410) has indicated that
Beryciformes differ from Perciformes in the number of infraorbitals and the extent
of the subocular shelf. He finds that in Beryciformes (except Holocentridae) there
are four circumorbitals behind the lachrymal whereas in Perciformes there are five,
and that in Beryciformes the subocular shelf extends over more than one infra-
orbital while in Perciformes it is confined to the second infraorbital. The subocular
shelf extends along all the infraorbitals in Holocentridae (living and fossil) and
in Polymixiidae (Polymixia, Homonotichthys) , but in Trachichthyidae (living
and fossil), Berycidae, Monocentridae, etc., the shelf is restricted to the second
infraorbital, as in Perciformes. The subocular shelf also extends along all the
infraorbitals in Anabantidae and Belontiidae (Liem 1963) : probably a subocular
shelf on all the infraorbitals, as in polymixiids, holocentrids and anabantids, is the
primitive condition of the structure (Smith & Bailey 1962 : 3). In the number of
infraorbitals the basic beryciform condition is undoubtedly as in percoids, with a
total of six bones, a lachrymal, four infraorbitals, and a dermosphenotic overlying
the autosphenotic : this condition occurs in Polymixiidae, living and fossil, and in
living Holocentridae. In Trachichthyidae, Berycidae, Monocentridae and some
Cretaceous Holocentridae (Caproberyx,Stichocentrus, Patterson 1964 : 347; 1967 : 89)
the apparent reduction in number of circumorbital bones is caused by fusion of
the dermo- and autosphenotics, which had already occurred in the Cretaceous
trachichthyid Hoplopteryx (Patterson 1964, fig. 55).
In summary, of the various beryciform-perciform lineages which have been
proposed, only the Aipichthys — carangid lineage emerges unscathed from a detailed
examination of caudal structures. The caudal skeletons of Polymixiidae, Sphen-
ocephalus, Dinopteryx and Pycnosteroides differ fundamentally from those of Perci-
formes in having a full neural spine on the second pre-ural centrum. In Aipichthys
and Pharmacichthys there are tendencies towards the Perciformes in the occasional
fusion of the second ural centrum with the preceding centrum and the occurrence
of five hypurals, but it is only among the Berycoidei that the basal perciform caudal
skeleton is duplicated. All known Berycoidei, even the very generalized Lissoberyx,
are more specialized than Perciformes in having procurrent caudal spines, and this
and characters of the skull (Patterson 1964 : 467) show that no perciform could
have evolved from any known berycoid. But if the Perciformes and cognate
groups (Channiformes, Scorpaeniformes, Pleuronectiformes, Tetraodontiformes)
should prove to be a monophyletic group, an alternative to the traditional method —
" an attempt first to define orders and other higher taxa and then to speculate upon
their origin, albeit in the light of the known fossils " (Greenwood et al. 1966 : 346)
is to use the criteria recommended by Hennig (1966 : 88, 120) and to search among
the living fauna for the sister group (Hennig 1966 : 139; see also Brundin 1966 : 17)
of this assemblage. A preliminary analysis suggests that the Berycoidei, not the
Beryciformes as a whole, may fill this role. This is indicated not only by features
of the caudal skeleton but by the fact that the Berycoidei is the only beryciform
group showing such perciform features as a subocular shelf confined to the second
infraorbital, the absence of epineurals, the pelvic girdle firmly joined to the cleithra
(in Berycidae especially), pelvic fins containing a spine and five soft rays (in Anomalo-
96 THE CAUDAL SKELETON IN
pidae and Gibberichthyidae, for example), partially separate soft and spinous dorsal
fins, etc., and in the stephanoberycoids, which appear to be merely specialized
offshoots of the trachichthyid lineage, loss of the orbitosphenoid. While it is clear
that many of these perciform characters in Berycoidei have arisen independently
within the group and were not inherited from a common ancestor of Berycoidei and
perciforms, they appear to be true parallelisms (Simpson 1961 : 78), and are
indicative of relationship.
(c) Intermediate groups.
Between the Beryciformes and the basal Percoidei, Greenwood et al. (1966 : 398)
place the Zeiformes, Lampridiformes, Gasterosteiformes, Channiformes, Synbranchi-
formes, Scorpaeniformes, Dactylopteriformes and Pegasiformes. Of the Gastero-
steiformes, Scorpaeniformes, Dactylopteriformes and Pegasiformes I have nothing
to say. The Channiformes (see Gosline 1968) and Synbranchiformes are probably
derived from the percoid level rather than from the beryciform or pre-beryciform.
In Zeiformes the caudal skeleton resembles those of basal Perciformes in having
no free second ural centrum and in having only five hypurals, and is more advanced
than basal percoids in having lost the second uroneural. In Zeidae (Zeus, Cyttus],
Oreosomatidae (Neocyttus) and Grammicolepidae (Xenolepidichthys] there is a com-
plete neural spine on the second pre-ural centrum, but this never occurs in conjunc-
tion with three epurals, and in Caproidae (Capros, Antigonia; Gosline 1961, fig. 4A),
which in other respects appear to have the most primitive caudal skeletons of the
group (the hypurals autogenous, the stegural autogenous in Antigonia) there are
three epurals and there is a low crest on the second pre-ural centrum, as in percoids.
Conditions in the Caproidae indicate that the neural spine on the second pre-ural
centrum in Zeidae, Oreosomatidae and Grammicolepidae has arisen secondarily by
fusion of the first epural (see above, p. 89, Fig. 28), and that as Gosline (1961) has
already said, there is nothing in the caudal skeleton to distinguish Zeiformes from
Perciformes. It has long been recognized that the Zeiformes are probably related
to the Beryciformes, principally because of the pelvic ray count, but they also show
many perciform features (Gosline 1961 : 36) and no conclusion has yet been reached
on whether they are more closely related to the beryciforms or the perciforms,
although in most recent classifications they are placed directly after the Beryci-
formes. Stinton (1967) has recently shown that there is a remarkable resemblance
between the otoliths of Antigonia and those of Berycoidei (Berycidae, Trachichthyi-
dae, and especially Monocentridae) . Stinton interprets this as indicating that
Antigonia is a berycoid, for he finds that the otoliths of Capros resemble those of
the zeids rather than Antigonia. However, the evidence that Antigonia and Capros
are related can hardly be ignored, and in Stinton's illustrations of zeiform otoliths
it seems possible to recognize a trend in reduction and specialization of the otolith
in the sequence Antigonia-Capros-Cyttus-Zeus: the same sequence of increasing
specialization is also shown by fusion within the caudal skeleton. In my opinion
Stinton's otolith evidence indicates not that Antigonia is a berycoid, but that as
the most primitive living zeiform it retains the clearest evidence of a common ancestry
with the Berycoidei. The percoid-like caudal skeleton, pelvic spine, etc., of the
MESOZOIC ACANTHOPTERYGIAN FISHES 97
Zeiformes do not oppose such a relationship. In Hennig's terminology, the Zei-
formes appear to be the apomorph sister group of the Berycoidei, these two groups
together forming the plesiomorph sister group of the perciform assemblage (see
above, p. 95).
The Lampridiformes, previously unknown before the Oligocene, have recently
acquired a respectable fossil record with Bonde's (1966) preliminary description of
a ? veliferid from the basal Eocene Mo-clay of Denmark, his opinion that Palaeo-
centrotus Kuhne (1941), from the same beds, is a lampridoid, not a zeiform, and the
suggestion (p. 81) that the Danian Bathysoma is a lampridiform, not a menid.
It appears that the deep-bodied Lampridiformes of the suborder Lampridoidei were
an important element of early Tertiary faunas. The caudal skeleton of Lampridi-
formes (known in Velifer, Lampris, Palaeocentrotus and Bathysoma) is characterized
by fusion of one or more of the upper hypurals with the second ural centrum, but
in other respects it does not differ from that of Beryciformes (there are six hypurals
in Velifer). In Lampris and Velifer the neural spine of the second pre-ural centrum
is reduced, but in Palaeocentrotus (Kiihne 1941, fig. 2) it is about half as long as its
predecessor, a primitive condition only found among Beryciformes in Aipichthys
(Fig. n) and Pharmacichthys. Further, in Lampris and Velifer there are seventeen
branched caudal rays and the caudal rays are deeply cleft basally, covering much of
the hypurals. These points tend to confirm the suggestion (Patterson 1964 : 473)
that the Lampridiformes are an offshoot of the Dinopterygoidei. Within the Dino-
pterygoidei, the second pre-ural neural spine, the seventeen branched principal
rays, the " hypurostegy " and the absence of a pelvic spine all point to the Aipich-
thyidae and Pharmacichthyidae as ancestral forms.
Transference of Bathysoma to the Lampridiformes leaves unsettled the position
of Mene, to which I thought Bathysoma was related (Patterson 1964 : 424). The
similarities between Mene and Bathysoma are numerous, and extend to the form of
the fin-rays, which are preserved in the middle part of the anal fin of the holotype
of B. lutkeni in Copenhagen, and are short, broad and unbranched (Bonde, personal
commn). The skull of Mene is also strikingly like that of Velifer (Regan 1907,
figs. 167, 169). But the caudal skeletons of Mene and the Lampridiformes are
very different. The six hypurals (in Velifer}, fusion of the second ural centrum with
the upper hypurals rather than with the preceding centrum, and the seventeen
branched principal rays of Lampridiformes can only be derived from beryciform or
pre-beryciform ancestors, but the caudal skeleton of Mene, with fifteen branched
principal rays and fusion of the second ural centrum and first four hypurals with the
preceding centrum could have evolved from the caudal skeleton of basal percoids
(p. 81). Nor can I find anything in the skull and vertebral column of Mene which
is against perciform ancestry. Nevertheless, Mene has an unusually long fossil
record, extending back to the Lower Palaeocene. The earliest recorded species,
M. phosphaticus Astre (1927) from the Montian of Tunisia, seems to agree with
Mene rather than with the Lampridiformes in caudal structure. Mene is very com-
mon in the Middle Eocene of Monte Bolca: with Mene at Monte Bolca there occur
other deep-bodied fishes of similar structure such as Exellia ( = Semiophorus) .
These fishes, though poorly known, seem to agree with Mene in the absence of spines
98 THE CAUDAL SKELETON IN
in the dorsal and anal fins (see Blot 1967 on Exellia} and also in the deeply cleft
bases of the caudal fin-rays. It is difficult to know how much significance can be
attached to this last character. E. & Y. Le Danois (1964), who coined the term
" hypurostegy " for it, give great importance to it and use it to unite in an " ordre
des Scombres " fishes as diverse as the holostean Pachycormidae, the Cretaceous
Tselfatiidae, the lampridiform Veliferidae and Lampridae, the carangids, scombrids
and others. In my opinion this assemblage is entirely spurious; the occurrence of
hypurostegy in such varied groups indicates not that the fishes are related but that
hypurostegy has arisen independently in a number of lines, for reasons as yet
unknown. E. & Y. Le Danois made a new family Vomeridae to include Mene, the
Eocene Vomeropsis, and the deep-bodied carangids Vomer, Selene, Alectis and
Hynnis, grouping this family with the Lampridae, Veliferidae, Ephippidae and
Exellia. Although the reasoning on which this grouping is made is doubtful, there
may be some truth in it. It seems possible that Mene and Exellia could represent an
independent attainment of the perciform grade from the Palaeocene lampridiform
stock. But the possibility that the resemblances between these Eocene forms and
the Lampridiformes are due to convergence is by no means ruled out: revisionary
studies on the Monte Bolca fauna now in progress (Blot 1967) may settle this question.
IV. CONCLUSIONS
Study of a single structural complex such as the caudal skeleton is unlikely to
produce firm conclusions on matters of phylogeny and relationships. Rather it
will serve as a means of checking existing hypotheses and will raise questions to be
settled by more comprehensive work. The main points arising from this paper are
as follows. An asterisk indicates that the genus or group is extinct.
1. Ctenothrissiformes,* Myctophoidei and Beryciformes have a basically similar
caudal skeleton, with the first ural and pre-ural centra fused, a free second ural
centrum, a stegural (the first uroneural fused with neural arch material from the
first ural and pre-ural centra), a second uroneural, three epurals and six hypurals.
The primitive nineteen principal caudal rays are retained in all ctenothrissiforms
and myctophoids, and in all beryciforms except the Polymixiidae, Sphenocephalidae,*
Dinopterygidae * and Pycnosteroididae,* which have eighteen.
2. Ctenothrissiformes * (Aulolepis, Ctenothrissa) and Myctophoidei (Aulopus,
Nematonotus,* Sardinioides* Acrognathus *) have a large caudal scute above and
below the caudal skeleton. The myctophoids Nematonotus * and Sardinioides
attenuatus * have a single urodermal on the base of the upper caudal rays. Both
caudal scutes and urodermals are relict structures absent in all higher groups.
3. The neural spine of the second pre-ural centrum in teleosts is primitively slender
and about half as long as its predecessor, as in Elops and Leptolepis.* This type of
second pre-ural neural spine persists in the myctophoids Aulopus and Nematonotus *
and in the beryciforms Aipichthys * and Pharmacichthys.* In Ctenothrissiformes,*
Ctenothrissa and Aulolepis have this short spine expanded into a plate, as it is in
most myctophoids, but in Pateroperca the spine is elongate and supports procurrent
fin-rays. A fully developed second pre-ural neural spine also occurs among Beryci-
formes in all polymixioids (Polymixiidae, Sphenocephalidae*) and in the dinop-
MESOZOIC ACANTHOPTERYGIAN FISHES 99
terygoids Dinopfayx* and Pycnosteroides.* In Berycoidei, as in generalized Perci-
formes, the second pre-ural neural arch and spine are reduced to a low crest.
4. The differences between the caudal skeletons of Myctophoidei and Cteno-
thrissiformes* and those of the most primitive teleosts are minor. Among primitive
teleosts the Clupavidae * come closest to the Ctenothrissiformes and Myctophoidei
in caudal anatomy.
5. The foremost procurrent caudal fin-rays are spinous in all living Beryciformes.
This is a feature peculiar to Beryciformes: Perciformes appear to be without
procurrent caudal spines. Procurrent caudal spines also occur in some species of
Ctenothrissa and in Aulolepis (Ctenothrissiformes*), probably an indication of rela-
tionship between Ctenothrissiformes and Beryciformes, although procurrent caudal
spines have also developed in advanced members of the myctophoid family Mycto-
phidae. Among Cretaceous Beryciformes, procurrent caudal spines occur only in
forms already known to be closely related to living Beryciformes (all Berycoidei and
the polymixiids Homonotichthys* and Pycnosterinx*) and in Dinopteryx.*
6. Within the Beryciformes, the principal variations in the caudal skeleton and
fin (apart from those in the second pre-ural neural spine, principal fin-ray count and
procurrent fin-rays already mentioned) are the presence of only two epurals in
Sphenocephalus* (Polymixioidei), the presence of only five hypurals in Aipichthys
velifer* some specimens of Pycnosteroides* (both Dinopterygoidei) and in many
Berycoidei (living holocentrids, Diretmus), and the fusion of the second ural centrum
with the preceding centrum in some specimens of Aipichthys* and in many Bery-
coidei (Berycidae, Diretmidae, Anoplogasteridae, living Holocentridae) . A full
neural spine on the second pre-ural centrum has been found in one individual of
Aipichthys* and one of Monocentris, simulating the polymixoid condition.
7. The basal perciform caudal skeleton differs from the basal beryciform type in
having only five hypurals, no free second ural centrum and only seventeen principal
rays. Perciformes are primitively characterized by having a low neural crest on
the second pre-ural centrum. Where a complete second pre-ural neural spine occurs
as a normal feature in perciform or higher groups the condition is secondary, the
spine representing an epural which has secondarily fused with the neural arch.
8. Evidence from the caudal skeleton does not support the various independent
beryciform — perciform lineages which have been proposed. The polymixioids and
the dinopterygoids Dinopteryx* and Pycnosteroides* differ fundamentally from the
various perciform groups which they otherwise resemble in having a fully developed
neural spine on the second pre-ural centrum. Pharmacichthys* which resembles
the acanthuroids and balistoids in many ways, differs from them in having the
bases of the caudal fin-rays deeply cleft, and is therefore unlikely to have been ances-
tral to these groups. Only the postulated link between Aipichthys* and the caran-
gids is not opposed by evidence from the caudal skeleton and fin.
9. Although no known berycoid could have been ancestral to any perciform, the
Berycoidei is the only beryciform group in which the caudal skeleton evolves towards
the percoid condition. There are many other characters and evolutionary trends
in which the Berycoidei is the only beryciform group to resemble the percoids : these
suggest that the Berycoidei is the sister group of the Perciformes and cognate groups.
TOO THE CAUDAL SKELETON IN
10. The caudal skeleton shows that the Danian Bathysoma* is a lampridiform
(the earliest yet known), not a member of the Menidae. It is suggested that the
Lampridiformes originated from near the beryciform families Aipichthyidae* and
Pharmacicthyidae.* In the Eocene there are fishes (Mene, Exellia*} which have
reached the perciform grade but resemble the Palaeocene and Eocene Lampridi-
formes : possibly these forms represent an independent attainment of the perciform
grade from lampridiform ancestors.
11. The caudal skeleton of the Zeiformes is basically of percoid type, but the
evidence of zeiform otoliths indicates that they are closely related to the Berycoidei.
The Zeiformes is evidently the sister group of the Berycoidei, these two groups
together being the sister group of the perciform assemblage.
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25 figs.
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BERRY, F. H. & ROBINS, C. R. 1967. Macristiella perlucens, a new clupeiform fish from the
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Monte Bolca (Italic). Collogues int. Cent. Natn. Rech. scient., Paris, 163: 133-138, i pi.
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BRUNDIN, L. 1966. Transantarctic relationships and their significance, as evidenced by
chironomid midges. K. svensha VetenskAhad. Handl., Stockholm, (4) 11, i : 1-472, 30 pis.
CAVENDER, T. 1966. The caudal skeleton of the Cretaceous teleosts Xiphactinus, Ichthyo-
dectes and Gillicus, and its bearing on their relationship with Chirocentrus. Occ. Pap. Mus.
Zool. Univ. Mich., Ann Arbor, 650 : 1-15, i pi.
DAVIS, J. W. 1887. The fossil fishes of the Chalk of Mount Lebanon, in Syria. Sci. Trans.
R. Dublin Soc., (2) 3 : 457-636, 25 pis.
1890. On the fossil fish of the Cretaceous formations of Scandinavia Sci. Trans. R.
Dublin Soc., (2) 4 : 367-434, 19 pis.
EJEL, F. & DUBERTRET, L. 1966. Sur 1'age precis du gisement de Poissons et de Crustaces
cretaces de Sahel Alma (Liban). C. r. Seanc. Soc. geol. Fr., Paris, 1966 : 353-354, i fig.
FRASER-BRUNNER, A. 1949. A Classification of the Fishes of the Family Myctophidae.
Proc. Zool. Soc. Lond., 118 : 1019-1106, 167 figs.
GOSLINE, W. A. 1960. Contributions toward a classification of modern isospondylous fishes.
Bull. BY. Mus. nat. Hist. (Zool.), London, 6 : 325-365, 15 figs.
1961. Some osteological features of modern lower teleostean fishes. Smithson. Misc.
Coll., Washington, 142, 3 : 1-42, 8 figs.
19610. The Perciform Caudal Skeleton. Copeia, Ann Arbor, 1961 : 265-270, 3 figs.
1963. Considerations regarding the relationships of the percopsiform, cyprinodontiform
and gadiform fishes. Occ. Pap. Mus. Zool. Univ. Mich., Ann Arbor, 629 : 1-38, 11 figs.
1965. Teleostean Phylogeny. Copeia, Washington, 1965 : 186-194, i fig.
1966. The limits of the fish family Serranidae, with notes on other lower percoids. Proc.
Calif. Acad. Sci., San Francisco, (4) 33 : 91-112, io figs.
MESOZOIC ACANTHOPTERYGIAN FISHES 101
GOSLINE, W. A. 19660. Comments on the Classification of the Percoid Fishes. Pacific Sci.,
Honolulu, 20 : 409-418, 2 figs.
1968. The Suborders of Perciform Fishes. Proc. U.S. Nat. Mus., Washington, 124:
1-77, 12 figs.
GOSLINE, W. A., MARSHALL, N. B. & MEAD, G. W. 1966. Order Iniomi. Characters and
synopsis of families. Mem. Sears Fdn Mar. Res., New Haven, 1, 5 : 1-18, 6 figs.
GREENWOOD, P. H. 1967. The caudal skeleton in osteoglossoid fishes. Ann. Mag. nat. Hist.,
London, (13) 9 : 581-597, 12 figs.
GREENWOOD, P. H., ROSEN, D. E., WEITZMAN, S. H. & MYERS, G. S. 1966. Phyletic studies
of Teleostean fishes, with a provisional classification of living forms. Bull. Amer. Mus.
Nat. Hist., New York, 131 : 339-456, pis. 21-23, 32 charts.
HAY, O. P. 1903. On a collection of Upper Cretaceous fishes from Mount Lebanon, Syria,
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Hist., New York, 19 : 394-452, 14 pis.
HENNIG, W. 1966. Phylogenetic Systematics. 263 pp., 69 figs. Translated by D. D. Davis
& R. Zangerl. Univ. of Illinois, Urbana.
HOLLISTER, G. 1937. Caudal skeleton of Bermuda Shallow Water Fishes. II. Order Per-
comorphi, Suborder Percesoces: Atherinidae, Mugilidae, Sphyraenidae. Zoologica, N.Y.,
22 : 265-280, 14 figs.
19370. Caudal Skeleton of Bermuda Shallow Water Fishes. III. Order Iniomi: Syno-
dontidae. Zoologica N.Y., 22 : 385-399, 1 8 figs.
HUSSAKOF, L. 1929. A new teleostean fish from the Niobrara of Kansas. Amer. Mus.
Novit., New York, 357 : 1-4, 2 figs.
KUHNE, W. G. 1941. A new Zeomorph Fish from the Paleocene Moler of Denmark. Ann.
Mag. nat. Hist., London, (n) 7 : 375-386, 3 figs.
LE DANOIS, E. & LE DANOIS, Y. 1964. L'Ordre des Scombres. Mem. Inst. fr. Afr. noire,
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LIEM, K. F. 1963. The comparative osteology and phylogeny of the Anabantoidei (Tele-
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1967. A morphological study of Luciocephalus pulcher, with notes on gular elements in
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MARSHALL, N. B. 1961. A young Macristium and the Ctenothrissid fishes. Bull. Br. Mus.
nat. Hist. (Zool.), London, 7 : 353-370, 4 figs.
MONOD, T. 1967. Le complexe urophore des Teleosteens: typologie et evolution (note pre-
liminaire). Colloques int. Cent. natn. Rech. scient., Paris, 163: 111-131, 16 figs.
NORDEN, C. R. 1961. Comparative osteology of representative Salmonid Fishes, with particu-
lar reference to the Grayling (Thymallus arcticus] and its Phylogeny. /. Fish. Res. Bd.
Can., Ottawa, 18 : 679-791, i6pls.
NURSALL, J. R. 1963. The hypurapophysis, an important element of the caudal skeleton.
Copeia, Ann Arbor, 1963 : 458-459.
NYBELIN, O. 1963. Zur Morphologie und Terminologie des Schwanzskelettes der Actinop-
terygier. Ark. Zool., Stockholm, (2) 15 : 485-516, 22 figs.
PATTERSON, C. 1964. A review of Mesozoic acanthopterygian fishes, with special reference
to those of the English Chalk. Phil. Trans. R. Soc., London, (B) 247 : 213-482, pis. 2-5.
1967. New Cretaceous berycoid fishes from the Lebanon. Bull. Br. Mus. nat. Hist.
(Geol.), London, 14 : 67-110, 4 pis.
19670. Are the teleosts a polyphyletic group? Colloques int. Cent. natn. Rech. scient.,
Paris, 163:93-109, n figs.
1968. The caudal skeleton in Lower Liassic pholidophorid fishes. Bull. Br. Mus. nat.
Hist. (Geol.), London, 16:201-239, 5 pis.
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stean fishes of the suborder Allotriognathi. Proc. Zool. Soc. Land., 1907 : 634-643, figs.
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102 THE CAUDAL SKELETON IN AC ANTHOPTERYGI AN FISHES
REGAN, C. TATE 1910. The caudal fin of the Elopidae and of some other teleostean fishes.
Ann. Mag. nat. Hist., London, (8) 6:354-358, 2 figs.
— 191 1. The anatomy and classification of the teleostean fishes of the Orders Berycomorphi
and Xenoberyces. Ann. Mag. nat. Hist., London, (8) 7 : 1-9, pi. i.
— 191 1«. The Anatomy and Classification of the Teleostean Fishes of the Order Iniomi.
Ann. Mag. nat. Hist., London, (8) 7 : 120-133, 7 figs.
ROSEN, D. E. 1962. Comments on the Relationships of the North American Cave Fishes of
the Family Amblyopsidae. Amer. Mus. Novit., New York, 2109 : 1-35, 24 figs.
1964. The relationships and taxonomic position of the halfbeaks, killifishes, silversides,
and their relatives. Bull. Amer. Mus. Nat. Hist., New York, 127 : 217-268, pis. 14, 15.
SIMPSON, G. G. 1961. Principles of Animal Taxonomy, xii + 247 pp., 30 figs. Columbia
Univ., New York.
SMITH C. L. & BAILEY, R. M. 1962. The Subocular Shelf of Fishes. /. Morph., Philadelphia,
110 : 1-18, 3 pis.
SMITH, J. L. B. 1951. The Fishes of the Family Veliferidae from South Africa. Ann. Mag.
nat. Hist., London, (12) 4 : 497-510, pis. 10-12.
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1962. Osteological studies on Pacific salmon of the genus Oncorhynchus . Bull. Fish.
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WEITZMAN, S. H. 1962. The osteology of Brycon meeki, a generalized characid fish, with an
osteological definition of the family. Stanford ichthyol. Bull., Palo Alto, 8 : 1-77, 21 figs.
1967. The origin of the stomiatoid fishes with comments on the classification of salmoni-
form fishes. Copeia, Washington, 1967 : 507-540, 18 figs.
WHITEHOUSE, R. H. 1910. The caudal fin of the Teleostomi. Proc. zool. Soc. Lond., 1910 :
590-627, pis. 47-50.
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Mag. nat. Hist., London, (n) 9 : 537-568, 5 pis.
VI. ABBREVIATIONS USED IN FIGURES
dz second, uroneural
e 1-3 epurals
f.s caudal scute
h 1-6 hypurals
hpu2 haemal spine of second pre-ural centrum
npu2, npu3 neural spines of second and third pre-ural centra
ph parhypural (haemal spine of first pre-ural centrum)
pui + ui centrum formed by fusion of first pre-ural and ural centra
puz, pus, pu4 second, third and fourth pre-ural centra
st stegural (first uroneural fused with pre-ural neural arch material)
U2 second ural centrum
Combinations of symbols linked by plus signs indicate compound elements formed by fusion
of the bones indicated.
PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING
NON-CALCAREOUS
MICROPLANKTON FROM THE
CENOMANIAN OF ENGLAND,
NORTHERN FRANCE AND
NORTH AMERICA
PART I
R. J. DAVEY
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 3
LONDON: 1969
NON-CALCAREOUS MICROPLANKTON FROM
THE CENOMANIAN OF ENGLAND,
NORTHERN FRANCE AND NORTH AMERICA
PART I
BY
ROGER JACK DAVEY
Pp. 103-180; ii Plates; 16 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 3
LONDON: 1969
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 17, No. 3 of the Geological
(Palaeontological] series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation :
Bull. Br. Mus. nat. Hist. (Geol.)
Trustees of the British Museum (Natural History) 1969
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 17 January, 1969 Price £2 i6s,
NON-CALCAREOUS MICROPLANKTON FROM
THE CENOMANIAN OF ENGLAND,
NORTHERN FRANCE AND NORTH AMERICA
PART I
By ROGER JACK DAVEY
Manuscript accepted May 1967
CONTENTS
SYNOPSIS
I. INTRODUCTION
Page
107
107
Acknowledgments . . . . . . . . no
II. STRATIGRAPHIC LOCATION OF SAMPLES . . . . . 112
Fetcham Mill Borehole . . . . . . . 112
Compton Bay ......... 112
Speeton .......... 112
Hunstanton ......... 115
Devon .......... 115
Escalles Borehole . . . . . . . . 115
Saskatchewan ......... 115
Texas .......... 115
III. SYSTEMATIC DESCRIPTIONS . . . . . . . 120
Genus Gonyaulacysta Deflandre . . . . . . 120
Gonyaulacysta cassidata (Eisenack & Cookson) . . . 120
whitei Sarjeant . . . . . . 120
fetchamensis Sarjeant . . . . . 120
exilicristata sp. nov. . . . . . 121
delicata sp. nov. . . . . . 123
sp. A. 124
Genus Cribroperidinium Neale & Sarjeant . . . . 125
Cribroperidinium intricatum sp. nov. . . . . 125
Other species ......... 128
Cribroperidinium orthoceras (Eisenack) . . . . 128
Genus Carpodinium Cookson & Eisenack . . . . 129
Carpodinium obliquicostatum Cookson & Hughes . . . 129
Genus Ellipsodinium Clarke & Verdier . . . . . 129
Ellipsodinium rugulosum Clarke & Verdier . . . . 130
Genus Apteodinium Eisenack . . . . . . 130
Apteodinium granulatum Eisenack . . . . . 130
Genus Trichodinium Eisenack & Cookson . . . . 131
Trichodinium castaneum (Deflandre) . . . . . 131
Genus Microdinium Cookson & Eisenack . . . . 132
Microdinium cf. ornatum Cookson & Eisenack . . . 132
setosum Sarjeant . . . . . . 133
distinctum sp. nov. . . . . . 133
variospinum sp. nov. . . . . . 135
veligerum (Deflandre) . . . . . 136
crinitum sp. nov. . . . . . 137
GEOL. 17, 3 i
106 CENOMANIAN NON-CALCAREOUS MICROPLAN KTON, i
Genus Histiocysta nov. . . . . . . . 138
Histiocysta palla sp. nov. . . . . . . . 138
Genus Fromea Cookson & Eisenack . . . . . 140
Fromea amphora Cookson & Eisenack . . . . 140
Genus Chytroeisphaeridia Sarjeant . . . . . 140
Chytroeisphaeridia euteiches sp. nov. ..... 141
Genus Cassiculosphaeridia nov. . . . . . . 141
Cassiculosphaeridia reticulata sp. nov. ..... 142
Genus Epelidosphaeridia nov. . . . . . . 142
Epelidosphaeridia spinosa (Cookson & Hughes) . . . 143
Genus Hystrichosphaeridium Deflandre . . . . . 143
Hystrichosphaeridium tubiferum (Ehrenberg) . . . 143
deanei Davey & Williams . . . 144
readei Davey & Williams . . . 144
radiculatum Davey & Williams . . 144
mantelli Davey & Williams . . 145
bowerbanki Davey & Williams . . 145
difficile Manum & Cookson . . . 145
Genus Oligosphaeridium Davey & Williams . . . . 146
Oligosphaeridium complex (White) . . . . . 146
reticulatum Davey & Williams . . . 147
prolixispinosum Davey & Williams . . 147
anthophorum (Cookson & Eisenack) . . 147
reniforme (Tasch) . . . . . 148
Genus Litosphaeridium Davey & Williams . . . . 148
Litosphaeridium siphoniphorum (Cookson & Eisenack) . . 148
Genus Polysphaeridium Davey & Williams . . . . 151
Polysphaeridium pumilum Davey & Williams . . . 151
laminaspinosum Davey & Williams . . 151
Genus Tanyosphaeridium Davey & Williams . . . . 151
Tanyosphaeridium variecalamum Davey & Williams . . 151
Genus Callaiosphaeridium Davey & Williams . . . . 151
Callaiosphaeridium asymmetricum (Deflandre & Courteville) . 152
Genus Cleistosphaeridium Davey, Downie, Sarjeant & Williams 152
Cleistosphaeridium heteracanthum (Deflandre & Cookson) . 152
multifurcatum (Deflandre) . . . 152
armatum (Deflandre) . . . . 153
polypes (Cookson & Eisenack) . . . 154
var. clavulum nov. . . . . . 154
huguonioti (Valensi) . . . . 155
var. pertusum nov. . . . . 156
flexuosum Davey, Downie, Sarjeant &
Williams ...... 157
parvum sp. nov. ..... 157
aciculare sp. nov. . . . . . 158
Genus Surculosphaeridium Davey, Downie, Sarjeant & Williams 158
Surculosphaeridium longifurcatum (Firtion) . . . . 158
Genus Hystrichokolpoma Klumpp . . . . . . 159
Hystrichokolpoma ferox (Deflandre) . . . . . 159
Genus Prolixosphaeridium Davey, Downie, Sarjeant & Williams 160
Prolixosphaeridium conulum sp. nov. . . . . . 160
Genus Coronifera Cookson & Eisenack ..... 161
Coronifera oceanica Cookson & Eisenack . . . . 162
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 107
Genus Exochosphaeridium Davey, Downie, Sarjeant & Williams 162
Exochosphaeridium phragmites Davey et al. . . . . 163
pseudohystrichodinium (Deflandre) . . 163
striolatum (Deflandre) . . . . 164
var. truncatum nov. . . . . 164
Other species ......... 166
Genus Cyclonephelium Deflandre & Cookson . . . . 166
Cyclonephelium distinction Deflandre & Cookson . . . 166
membraniphorum Cookson & Eisenack . . 167
vannophorum sp. nov. . . . . 168
paucispinum sp. nov. . . . . 170
eisenacki sp. nov. . . . . . 170
Genus Adnatosphaeridium Williams & Downie . . . 171
Adnatosphaeridium chonetum (Cookson & Eisenack) . . 171
Genus Hystrichosphaera O. Wetzel . . . . . . 172
Hystrichosphaera ramosa (Ehrenberg) . . . . . 172
var. ramosa (Ehrenberg) . . . . 172
var. gracilis Davey & Williams . . . 172
var. multibrevis Davey & Williams . . 173
var. reticulata Davey & Williams . . 173
cingulata (O. Wetzel) . . . . 173
var. reticulata Davey & Williams . . 174
crassimurata Davey & Williams . . . 174
crassipellis Deflandre & Cookson . . 174
Genus Achomosphaera Evitt. . . . . . . 174
Achomosphaera ramulifera (Deflandre) . . . . 174
sagena Davey & Williams . . . . 174
Genus Hystrichodinium Deflandre . . . . . . 174
Hystrichodinium voigti (Alberti) . . . . . . 175
dasys sp. nov. . . . . . . 175
IV. REFERENCES . . . . . . . . . .176
SYNOPSIS
This paper, which will appear in two parts, presents the results of a detailed study of some
non-calcareous microplankton from the Upper Cretaceous, and in particular of assemblages of
Cenomanian age. The stratigraphical potentialities of the fossil microplankton are briefly
assessed by the analysis of samples from five localities in England and one in France. To
assess the potentialities of long-range correlation, assemblages from Saskatchewan and Texas
have also been examined. Both quantitative and qualitative methods have been employed
and the correlations, both intra- and inter-regional, are promising. Seven new genera and
thirty-five new species and varieties are described.
I. INTRODUCTION
FOSSIL non-calcareous microplankton consist mainly of cysts of dinoflagellates,
together with various forms of unknown affinity placed in the Group Acritarcha
Evitt (1963). The majority of dinoflagellates are free-living, oceanic and plank-
tonic. They have a complex life-cycle, usually composed of four stages, in one of
which (the motile stage) they are capable of limited vertical movement by the use
of two flagella. During the life cycle, if the organism is subject to adverse con-
ditions, a resting cyst is formed. This, most palynologists believe, is the only stage
io8
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
in the dinoflagellate life-cycle preserved in the fossil state. Fossil non-calcareous
microplankton are useful as stratigraphic indices because they are planktonic, of
relatively resistant composition, abundant in most marine sedimentary samples, and
easy to extract. Hence the principal object of the study was to assess how accurate
dinoflagellate cysts are for intra- and inter-regional stratigraphic correlations.
The order of description of the dinoflagellate cyst-families follows that in Sarjeant
FIGURE ONE
MAP of SPECIMEN LOCALITIES
in ENGLAND and FRANCE
FIG. i. Map of specimen Localities in England and France.
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 109
& Downie (1966). The adjectives used when describing the frequency of a species
in an assemblage have been defined as follows: —
Very Common 10-100%
Common 1-10%
Infrequent 0-1-1%
Rare 0-01-0-1%
Very Rare under o • o i %
The first fossil microplankton were described and figured by Ehrenberg (1838,
1843, 1854) and included forms embedded in flakes of Upper Cretaceous flint from
Germany and Denmark. These observations stimulated English microscopists in
the mid-nineteenth century into finding these organisms in English flints of the same
age.
Little further research in this sphere was published until 1933 when O. Wetzel
described assemblages from Upper Chalk flints of the Baltic region. In 1934
Deflandre published the first of a number of well illustrated papers dealing with
fossil microplankton from the flints of the Paris Basin. Unfortunately the strati-
graphic horizons of the flints are unknown, many being picked up in the streets of
Paris. His two most important papers describing Upper Cretaceous microplankton
were published in 1936 and 1937. They contain accurate figures and descriptions
of many new species and genera mainly from the Senonian, although some of the
flints are probably of Cenomanian and Turonian age.
Between 1936 and 1952 few papers dealing with microplankton from the Upper
Cretaceous were published. Firtion (1952) described the first definite Cenomanian
assemblage, the material having been obtained from the Lower Cenomanian of
France. All his species have subsequently been observed in the Lower Chalk of
England and France except for Pareodina sp. which is unidentifiable. Firtion's
Hystrichosphaeridium cf . salpingophorum may correspond to H. mantelli, and Micrhy-
stridium ambiguum is probably Cleistosphaeridium huguonioti.
Since 1955 a number of publications have been produced dealing mainly with the
systematics of Cretaceous microplankton. Assemblages have been described from
Germany by Gocht (1957, 1959), Eisenack (1958), and Alberti (1959, 1961), but only
the latter author records Cenomanian species. Alberti (1961) described a series of
assemblages from the Valanginian to Turonian of northern Germany. Eight species
were recorded from the Cenomanian and all of these, except for Korojonia dubiosa,
have been found during the present study. Hystrichodinium pulchrum, identified by
Alberti from the Cenomanian, is here included within Hystrichodinium voigti. Gony-
aulax orthoceras, figured by Alberti, is undoubtedly Cribroperidinium intricatum sp.
nov., and Palaeohystrichophora cf. paucisetosa is probably a form of P. infusorioides
possessing fewer spines than usual.
Lower Cretaceous assemblages were described by Neale & Sarjeant (1962), Pocock
(1962), and Tasch, McClure & Oftedahl (1964). Gorka (1963) described nine species
of microplankton from the Cenomanian of Poland. All of these species have been
found in the Cenomanian deposits of England and France, although some are con-
sidered to be incorrectly identified. Gonyaulax orthoceras, illustrated by Gorka
no CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
appears to be Gonyaulacysta exilicristata sp. nov. and her Gonyaulax sp. is probably
Cribroperidinium intricatum ; Hystrichosphaeridium asterigerum is probably equivalent
to Oligosphaeridium complex and Hystrichosphaeridium polytrichum possibly to
Cleistosphaeridium armatum. Her Hystrichosphaeridium cf. striolatum may also
belong to the latter species.
Baltes (1963) described eight species of microplankton from the Cenomanian
deposits of Roumania. Of these seven have been recorded in the present study.
The species not recorded, Ascodinium hialinium, probably belongs to the genus
Deflandrea. The identities of three other species are in doubt: some of the specimens
illustrated as Hystrichosphaeridium longifurcatum probably belong to this species
(transferred to Surculosphaeridium by Davey et al. 1966), but one specimen (pi. 7,
fig. 12) probably belongs in Hystrichosphaera. Histrichosphaeridium sp. 22 resembles
Exochosphaeridium striolatum var. truncatum nov. and Hystrichosphaeridium sp. 23 is
possibly Cleistosphaeridium multifurcatum.
Manum & Cookson (1964) describe species of supposed lower Upper Cretaceous age
from Arctic Canada and of these, eight have also been recorded from the Lower
Chalk of England and France; however, the Arctic Canadian assemblages are more
comparable to those obtained from Saskatchewan.
Cookson & Hughes (1964) gave the first account of microplankton from the
deposits of Upper Albian and basal Cenomanian age in England. Thirty- three species
were described from the Cenomanian and of these only six have not been recorded
from the basal Cenomanian of Fetcham Mill (sample FM 840) and Compton Bay
(CB i).
A number of papers have been published (between 1954 and 1965) dealing with the
microplankton of Australia and, in part, of New Guinea and Papua — Deflandre &
Cookson (1954, 1955), Cookson (1956, 1965), Cookson & Eisenack (1958, 19600, b,
1961, 19620, b) Eisenack & Cookson (1960), and Cookson & Manum (1964). Un-
fortunately it is usually only possible to give the approximate age of the samples
and, therefore, stratigraphic conclusions are not as meaningful as one would have
hoped.
A number of species from the Cenomanian deposits of England were described by
Davey, Downie, Sarjeant & Williams (1966) and the present paper published in
two parts forms a natural continuation of that work.
Clarke & Verdier (1967) describe microplankton assemblages of Cenomanian to
Senonian age from the Isle of Wight, southern England. The Cenomanian samples
examined by them were obtained from a locality approximately sixteen miles to the
east of Compton Bay. Their results indicated that the Upper Cretaceous could be
divided into 5 zones and 5 subzones, and also into 7 " intervals " based on extinction
points. The results concerning the Cenomanian are mainly substantiated in the
present study.
ACKNOWLEDGMENTS
The research has been carried out during the tenure of a research studentship in the
Department of Geology, The University of Nottingham.
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
646' 1
0)
1
s
0
-8
0)
_Q
72 7'*
<D
5
N
,0
h.
0
84l'l
XX
Turonian
plenus Marls
XX
i i
•650-51
- A 7O - 71X
Ligh
Ch(
/
Grey
Ch«
QIC7
r grey
alk
^
marly
alk
i i
i i
i i
i i
i- 690-91
- 710-lf
i i
i
i i
i i
i
}"
- 7*5 f\ *5 1
/oU o I
- 750-5 f
-770-71'
i
i
i i
i
i
i
r 790-91
h810-llx
i
•815 -
Grey -green Marl
OO £.'
i
~ 836 -
'Chloritic'Marl
I
i i
fj^LlJ *\ \
Albion Upper Greensand
FIG. 2. Section showing the Location of the Samples analysed from
Fetcham Mill, Surrey. Scale i inch to 30 feet.
H2 CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i
The author would like to thank sincerely Dr. W. A. S. Sarjeant for considerable
help and encouragement given at all stages during the course of this work, and also
particularly acknowledges the interest and encouragement given by Professor W. D.
Evans and Dr. A. J. Rowell. Thanks are due to Professor G. Deflandrefor courtesy
in entertaining the author and allowing him to examine type material at the Labora-
toire de Micropaleontologie, Ecole Practique des Hautes Etudes, Paris ; and to Dr.
G. L. Williams for his collaboration when dealing with the systematics of certain
genera. The author is indebted to Mr. R. Hendry and his staff in the Department
of Geology, The University of Nottingham for their assistance and provision of
necessary laboratory equipment.
Special thanks are due to Sir James Stubblefield, former Director of the Institute
of Geological Sciences, for permitting study of chalk samples from the Fetcham
Mill Borehole; to the Bureau de Recherches Geologiques et Minieres, for providing
chalk samples from the Escalles Borehole ; to the Department of Mineral Resources,
Saskatchewan, for providing Cretaceous samples from the International Yarbo
Borehole no. 17; to Dr. J. D. Powell, for providing Upper Cretaceous samples from
Texas; to Dr. W. E. Smith, for providing samples from the Cenomanian of south
Devon; and to Dr. K. Diebel, of the Institut fur Palaeontologie, Humboldt Uni-
versity, East Berlin, for courteously permitting the loan of Ehrenberg's holotypes.
The following abbreviations are used in the text: B.M. (N.H.) — British Museum
(Natural History) : G.S.M. — Institute of Geological Sciences, London.
II. STRATIGRAPHIC LOCATION OF SAMPLES
1. Fetcham Mill Borehole, Leatherhead, Surrey (TQ 15815650).
This borehole has been described by Gray (1965). Ten samples at 20-foot intervals
were processed from the Cenomanian (Lower Chalk) succession (Fig. 2). In addition
one sample from the Albian (Upper Greensand) and one from the Turonian (Middle
Chalk) were analysed for comparative purposes.
2. Compton Bay, Isle of Wight (SZ 365854)
The Cenomanian is well exposed in the cliff section at this locality and has been
described by Jukes Browne (1903) and Osborne White (1921). Samples were
collected at 7-8 ft. intervals and eleven samples, at approximately 14 ft. intervals,
were analysed for their organic-shelled microplankton content (Fig. 3).
3. Speeton, Yorkshire (TA/i6675o)
The Cenomanian succession is fully exposed in the cliffs at Speeton. The base of
this stage was taken to coincide with the bottom of bed V (Wright 1963) which,
together with the overlying bed U, is placed in the Red Chalk (Fig. 4). The succes-
sion has been described recently by Kaye (1964). Seven samples were collected
and analysed for their microplankton content.
CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i
1.5 9'
Turanian plenus Marls
> i
^ subglobosus Zone '
i
•151-CB21
- 137- 19
-116- 17
- 107- 15
-92- 13
• 76'- 11
Ligh
Ch
/
Gre)
C
t grey
alk
t
/ marly
:halk
l
i
l
l
ES
l
I
I
I
l
I
l
I
i
i
1
l
I
l
1
1
i
i
l
l
1
I
l
i
1
1
1
i
i
\ varians Zone
- 59 - 9
h44- 7
- 28'- 5
• 15'- 3
l
l
1
I
l
I
l
i
l
1
l l
1 1
i i
i i
i i
•^_ Phosphatic nodules
i i
I i
i i
Blue-grey Chalk
.11' A"
i i
i i
i l
Chloritic Marl
i i
l l
V
' Albion
23'
21' 6"
FIG. 3. Section showing the Location of the Samples analysed from
Compton Bay, Isle of Wight. Scale i inch to 20 feet.
114 CENOMANIAN NON-CALCAREOUS MICROPLAN KTON, i
122-
Zone
subg
**
78'.
varians Zone
Turanian
T , I
II
^1
II
TTI
"115-Sp7
100- 6
78- 5
52'- 4
28-3
•12'- 2
0' - 1
plenus Marls
Grey -white
Chalk
- 80
Hard grey Chalk
-. 78'
Grey Chalk
8'
Marly Chalk
•28'
Pink Chalk U
Green -white Chalk V
Aibian
FIG. 4. Section showing the Location of the Samples analysed from
Speeton, Yorkshire. Scale i inch to 20 feet.
CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i 115
4. Hunstanton, Norfolk. (TF 675420)
The varians and subglobosus zones are exposed in the Hunstanton cliffs (Peake &
Hancock 1961). Three samples were processed, two from the lower zone and one
from the base of the Totternhoe Stone (Fig. 5).
5. Devon, South Coast
The Cenomanian is represented between Salcombe and Lyme Regis by isolated
patches composed of a few feet of sandy Cenomanian Limestone. These deposits
were divided into four beds by Jukes-Browne (1903) — Ar, A2, B and C. Bed C is
probably the Actinocamax plenus Marls. The samples were collected from four
localities by Dr. W. E. Smith (Fig. 6) :
(i) Maynards Cliff (see Smith 1961 : 114)
(ii) Beer Head (see Smith 1957 : 123)
(iii) Whitediff (see Smith 1957 : 118)
(iv) Humble Point (see Smith 1965 : 126)
6. Escalles Borehole, Cap Blanc-Nez, Pas de Calais
The borehole, drilled by the Bureau de Recherches Geologiques et Minieres in 1958
has been described by Destombes (1961). Eleven samples were obtained for analysis
at about 20 ft. (6m.) intervals (Fig. 7).
7. Saskatchewan, south-east
The borehole, from which the Saskatchewan samples were obtained, was drilled
for the Department of Mineral Resources, Saskatchewan and is called " International
Yarbo, no. 17 ". It is located east of Regina at Lsd. i, Sec. 24, Twp. 20, Rg. 33,
Wrst Meridian. All depths are measured from the Kelly Bushing which is at an
elevation of 1,690 ft. above sea level. Six samples of Albian/Cenomanian age were
analysed for their microplankton content (Fig. 8).
8. Texas, north
Two samples were obtained from the Upper Cenomanian of north Texas (Tarrant
County) for the author by Dr. J. D. Powell. The lower sample (T5) was obtained
from the Upper Woodbine Formation (Acanthoceras wintoni Zone) and consists of a
yellowish, slightly calcareous clay. The higher sample (T4) is from 35 ft. above the
base of the Eagle Ford Formation (Eucalycoceras Zone). This is a thin-bedded
yellow limestone containing shelly fossils and plant debris, in particular leaf
fragments.
n6 CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i
r20'
JS
C
o
N
-13' 10" - H2
•18' 6" - H3
HI
-18'
Totternhoe Stone
Creamy -white
Chalk
4 1
1 5
Inoceramus Bed
Paradoxica Bed
Red Rock
FIG. 5. Section showing the Location of the Samples analysed from
Hunstanton, Norfolk. Scale i inch to 2 £ feet.
CENOMANIAN NON-CALCAREOUS MICROPLAN KTON, i
117
n8
CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i
502
N
«/>
2
o
i
60f-
0)
c
O
M
716
Turonian
' - E153
-521' 6"
- 159
-541 - 165
-561' - 171
-5806- 177
-600' - 183
•620 - 189
639V- 195
659'6"- 201
679' - 207
698' 6'- 213
Grey- white
Chalk
Grey marly
Chalk
Chloritic Marl
712' 6*
Albian
FIG. 7. Section showing the Location of the Samples analysed from
Escalles, France. Scale i inch to 30 feet.
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 119
Q_
ID
o
0£
O
o
o
O
u
1
tt
Shale
/
Sas
UJ
Q_
Q.
ID
Second White Speckled Shale
007'
-835V
-QO/
-890'
Shale
c960
ex.
\* f \j \J
-967'
UJ
Fish Scale Zone
o
-c1030'
-1023
Shale
~l-IVJ«jw
-1084'
Viking
FIG. 8. Section showing the Location of the Samples analysed from
Saskatchewan, Canada. Scale i inch to 50 feet.
GEOL. 17, 3
120 CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
III. SYSTEMATIC DESCRIPTIONS
Class DINOPHYCEAE Pasher
Subclass DINIFEROPHYCIDAE Bergh
Cyst-FamUy GONYAULACYSTACEAE Sarjeant & Downie 1966
Genus GONYAULACYSTA Deflandre emend. Sarjeant 1966
Gonyaulacysta cassidata (Eisenack & Cookson) emend. Sarjeant
1960 Gonyaulax helicoidea subsp. cassidata Eisenack & Cookson : 3, pi. i, figs. 5, 6.
ig66a Gonyaulacysta cassidata (Eisenack & Cookson) Sarjeant : 125, pi. 14, figs. 3, 4, text-fig. 31
(see also for earlier references).
1967 Gonyaulacysta cassidata (Eisenack & Cookson) Clark & Verdier: 29, pi. 4, figs. 4-6.
DIMENSIONS. Range of observed specimens: overall length 59 (67-5) 78/4, overall
width 40 (46-4) 60 fi. Number of specimens measured, 14.
REMARKS. The Cenomanian specimens examined are very similar to the Austra-
lian Aptian-Cenomanian forms of Eisenack & Cookson (1960) and Cookson & Eisenack
(19626) except that the former are slightly smaller in size.
OCCURRENCE. G. cassidata is an infrequent species at all horizons throughout
the Cenomanian of Fetcham Mill, Compton Bay and Escalles. It has not been
recorded in the North American samples.
Gonyaulacysta whitei Sarjeant
ig66a Gonyaulacysta whitei Sarjeant : 126, pi. 14, fig. 2, text-fig. 32.
DIMENSIONS. Range of observed specimens: overall length 55-62/4, overall
width 42-50/4. Number of specimens measured, 5.
REMARKS. The shape of the shell, the tabulation and the crests make G. whitei
an easily recognizable and distinctive species. G. cf. ambigua Cookson & Eisenack
(19606) from the Upper Jurassic is of the same shape and possesses similar crests and
tabulation; plate i"" is absent. However, the apical horn of G. cf. ambigua is
rudimentary or absent, thus making differentiation easy.
OCCURRENCE. Of five specimens of G. whitei observed, four are from sample FM
770 and one from sample FM 750. The restricted distribution of this species and its
similarity to G. cf . ambigua from the Upper Jurassic suggest the possibility that this
is a derived form.
Gonyaulacysta fetchamensis Sarjeant
19660 Gonyaulacysta fetchamensis Sarjeant : 128, pi. 15, figs, i, 2, text-fig. 33.
REMARKS. G. fetchamensis has a rather unusual tabulation — two posterior inter-
calary plates and a seventh postcingular plate — and, as pointed out by Sarjeant
(19660), may subsequently form the basis of a new genus. As yet only two specimens
have been studied and this species has, therefore, been placed in Gonyaulacysta. It
CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i 121
appears to be a transitional type from the normal Gonyaulacysta to forms now placed
in Cribroperidinium Neale & Sarjeant. The latter forms are of similar overall appear-
ance but the shell is divided by low crests into an unusually large number of areas.
OCCURRENCE. G. fetchamensis is a rare species recorded only from the Chalk of
H.M. Geological Survey borehole, Fetcham Mill, Surrey, at 840 ft. depth. Upper
Cretaceous (Lower Cenomanian).
Gonyaulacysta exilicristata sp. nov.
(PI. i, figs, i, 2; Figs. gA, B)
DERIVATION OF NAME. Latin, exilis, thin or poor; cristatus, crested — with re-
ference to the poorly defined sutural crests.
DIAGNOSIS. Shell subspherical ; moderately well developed apical horn. Shell
wall thick, finely but densely granular. Reflected tabulation 3', la, 6", 6c, 6'"
(—7"'?), i p, i"". Plate boundaries marked by low, poorly defined crests which
sometimes form small spines at crestal nodes. Cingulum narrow, weakly laevo-
rotatory; sulcus of moderate width, widening slightly posteriorly.
HOLOTYPE. G.S.M. slide PF 3987 (i). Lower Chalk, H.M. Geological Survey
Borehole, Fetcham Mill, Surrey at 730 ft. depth. Upper Cretaceous (Cenomanian).
DIMENSIONS. Holotype: overall length 83/4, overall width 68 ju,, length of horn
I2/*. Range: overall length 70 (81-6) 98^, overall width 58 (64-5) 71/4. Number
of specimens measured, 24.
DESCRIPTION. The shell wall measures between 2 and 3/4 in thickness, the
endophragm being approximately twice as thick as the periphragm. The latter
forms the apical horn which is triangular in cross-section due to the sutural ridges
which delimit the apical plates extending along it. Lines of ornamentation, some-
times similar to the sutural crests, are present on some of the plates, particularly
those in the postcingular series. Occasionally it appears that plate 4'" is subdivided
by a low ridge so giving seven postcingular plates.
The cingulum is fairly narrow (3 to 47*) and only slightly laevo-rotatory. The
sulcus often possesses a posterior ventral plate. In the medial-posterior part of the
furrow there is usually an elongate depression, as seen in the holotype (Fig. gA). A
precingular archaeopyle is typically present.
REMARKS. G. exilicristata sp. nov. is distinguished from all previously described
species by its overall shape, the type of plate boundaries and the tabulation.
Apteodinium granulatum Eisenack (1958) is similar but has a stouter apical horn
and a tabulation appears to be absent; the cingulum is only rarely visible.
Cribroperidinium orthoceras (Eisenack) comb. nov. is also similar but possesses a
longer apical horn, and the tabulation differs and is more clearly defined.
OCCURRENCE G. exilicristata is a rare species recorded from a number of horizons
throughout the Cenomanian of Fetcham Mill, Compton Bay and Escalles. At only
one horizon is this species common, in sample FM 730 from Fetcham Mill. Two
specimens have been recorded from Saskatchewan, both from sample Sas 1084.
122 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
FIG. 9. Gonyaulacysta exilicristata sp. nov., A. Ventral Surface of Holotype (X 900),
B. Dorsal Surface of Holotype, (X 900). Gonyaulacysta Sp. A., C. Ventral Surface slide
PF. 3987 (2) (X 900). D. Dorsal Surface slide PF. 3987 (2) (X 900)
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
123
Gonyaulacysta delicata sp. nov.
(PI. i, figs. 7, 8; Figs. loA, B)
DERIVATION OF NAME. Latin, delicatus, delicate — with reference to the delicate
nature of the shell.
DIAGNOSIS. Shell subspherical, epitract and hypotract of similar size. Shell
wall thin, smooth. Apical horn absent, there being a small circular apical plate in
this position surrounded by three large apical plates. Reflected tabulation 4', la,
6", 6c, 6"', ip, i"". Plate boundaries well defined by low crests. Cingulum wide,
strongly laevo-rotatory ; sulcus broad.
HOLOTYPE. B.M. (N.H.) V. 51979(1). Lower Colorado, Second White Speckled
Shale, International Yarbo Borehole No. 17, Saskatchewan at 835 ft. depth.
Upper Cretaceous (Cenomanian).
PARATYPE. B.M. (N.H.) V. 51979(2).
DIMENSIONS. Holotype: length of shell 57/4, width 51/4. Paratype: length of
shell 55/4, width 47/4. Range: length of shell 52-60/4, width 47-51/4. Number of
specimens measured, 4.
DESCRIPTION. The shell wall is very thin (less than 0-5 ju, thick) and only attains
a thickness of 0-5 /* when forming the plate boundaries. Due to the thinness
of the shell wall specimens are easily distorted. The tabulation has, however, been
formulated after the examination of a number of specimens.
A B
FIG. 10. Gonyaulacysta delicata sp. nov., A. Ventral surface of Holotype (x 15°°)-
B. Dorsal surface of Holotype (X 1500).
124 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
Anteriorly three apical plates (2', 3' and 4') abutt against a small circular plate,
plate i', which is in the position normally occupied by the apical horn. The pre-
cingular and postcingular series of plates are large and more or less pentagonal.
Plate 3" is always lost in archaeopyle formation. The first postcingular plate is
elongate and has a poorly defined sulcal border.
The cingulum is wide (4 to 7 fi) and tends to be constricted at the cingular plate
boundaries. The sulcus is broad and widens slightly towards the posterior. The
anterior end of the sulcus has a flat margin from which arise plates i' and la. At
the posterior end of the sulcus there may be developed a posterior ventral plate.
REMARKS. The presence of very low sutural crests, the lack of an apical horn and
the tabulation differentiate G. delicata sp. nov. from all previously described species.
G. ambigua Deflandre, from the Kimeridgian of France, is of similar appearance
but possesses a small apical horn and differs in tabulation detail.
OCCURRENCE. G. delicata has been found in only one sample, Sas 835, and it is
there infrequent. This restricted distribution may well indicate that this is a
derived species.
Gonyaulacysta sp. A.
(PL i, figs. 9, 10; Figs. gC, D)
DESCRIPTION. Only one well preserved specimen of this species has so far been
observed. It possesses a subspherical shell, bearing a moderately well developed
horn with a trifid termination. The shell wall (c. 2/1 thick) is irregularly studded
with granules of varying shapes and sizes. The sutural crests are quite well defined,
but low, and indicate a reflected tabulation of 3', la, 6", 6c, 6'" (—7'"?), ip., i"".
The crests, demarcating the three apical plates, extend along the apical horn and
give rise to three small spines at its distal termination. Plates 2", 3" and 4" are
relatively large, plates i" and 5" rather elongate and plate 6" is reduced due to the
anterior intercalary plate. In the postcingular series, plates i"' and 2'" are
reduced and plate 7'" does not have a clearly marked plate boundary. Plate 2'"
possesses a curved line of ornamentation which is characteristic of Cnbroperidinium
Neale & Sarjeant. There is a single posterior intercalary plate and a large antapical
plate.
The cingulum is strongly laevo-rotatory and varies considerably in width (2-5 (JL),
being constricted at the cingular plate boundaries and expanding on either side.
The sulcus is broad and possesses a central depressed area of elongate shape. A
archaeopyle is present.
FIGURED SPECIMEN. G.S.M. slide PF. 3987, specimen 2. Lower Chalk, H.M.
Geological Survey Borehole, Fetcham Mill, Surrey at 730 feet depth, Upper Cretaceous
(Cenomanian) .
DIMENSIONS. Overall length 75/11, overall width 65^, length of horn 14 p..
REMARKS. The form of the apical horn, the type of granulation and the shape
of the cingular plates distinguish this species from all described forms. The shape
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 125
of the shell and the tabulation are most comparable to G. exilicristata suggesting a
relationship between the two species.
Genus CRIBROPERIDINIUM Neale & Sarjeant emend.
EMENDED DIAGNOSIS. Proximate cysts, subspherical to ovoidal, thick-walled.
Crests numerous and well developed. Tabulation ?6', (i-5a), 8-9", oc, g"', ip,
i-3p.v. (5-7 p.c.), o"" (— ?2""). Cingulum laevo-rotatory. Archaeopyle
precingular.
TYPE SPECIES. Cribroperidinium sepimentum Neale & Sarjeant 1962.
REMARKS. The diagnosis has been emended to draw attention to the fact that
the anterior intercalary series and the posterior circle series of plates are not always
readily distinguishable. The tabulation attributed to this genus by Neale &
Sarjeant was 6', i~5a, 8", 9'", ip, i-2p.v. 6-Pyp.c., o"". In determining the tabula-
tion they numbered all the delimited areas on the shell surface in the normal manner.
This numbering procedure has been used in the study of C. intricatum sp. nov. with
certain reservations. Difficulties arose due to the large number of delimited areas
and also to some variation on the dorsal surface of the hypotract. Some of the
crests are rudimentary and others, although appearing fairly normal, are unusual
in their position. It was discovered that by the removal of these unusual crests a
normal Gonyaulax-type tabulation could be reconstructed. This is clearly shown in
Figs. nA, B, of the ventral surface of the holotype of C. intricatum. Thus Cribro-
peridinium is basically a form of Gonyaulacysta which possesses additional crests.
These additional crests may correspond to an increase in the number of thecal
plates composing the motile dinoflagellate, but this is thought to be unlikely since:
(a) the plates formed would be of an extremely unusual shape ;
(b) these crests subdivide detached opercula demonstrably composed of a single
precingular plate (3") ;
(c) many of these crests are poorly developed and show considerable positional
variation on the dorsal surface of the hypotract; and
(d) that by their removal a normal Gonyaulax-type tabulation remains. Thus,
these additional crests may be regarded as an ornamentation or perhaps a rather,
superficial cyst strengthening device. It is, therefore, considered more practical
and correct to use a different numbering system for the Cribroperidinium cyst
tabulation, vis, roman numerals.
Three species, Gonyaulacysta orthoceras (Eisenack), G. muderongensis (Cookson &
Eisenack) and G. edwardsi (Cookson & Eisenack), are here transferred to Cribroperi-
dinium. This genus appears to be very limited in stratigraphic range (Hauterivian—
Lower Turanian) and, as such, is a useful stratigraphic indicator.
Cribroperidinium intricatum sp. nov.
(PL 2, figs. 1-3; Figs, n, 12)
DERIVATION OF NAME. Latin, intricatus, complicated — with reference to the
complex crest arrangement.
126
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
DIAGNOSIS. Shell subspherical, epitract and hypotract of similar size. Apical
horn of moderate length, subconical. Shell wall moderately thick, granular and
bears a few, randomly arranged tubercles. Crests usually in form of low ridges, well
denned, sometimes membranous, delimiting a large number of plates on shell
surface. Sutural spines absent. Plate IF" crossed diagonally by low crest.
Operculum possessing semi-circular crest. Sulcus possessing posterior ventral plates.
Cingulum narrow, plates not defined.
HOLOTYPE. B.M. (N.H.). V. 51980 (i). Upper Lower Colorado, Fish Scale Zone,
International Yarbo Borehole No. 17, Saskatchewan at 1,023 feet depth. Lower
Cretaceous (Albian).
DIMENSIONS. Holotype: overall length 120 /*, overall width 114/1,, length of horn
20 p. Range: overall length 107 (125-1) 142/11, overall width 101 (108-2) 126/1..
Number of specimens measured, 17.
DESCRIPTION. The shell wall is i to 1-5 p in thickness and densely granular. The
crests are typically low thickenings of the periphragm (2-3 p wide), but in the ant-
apical region, and occasionally elsewhere, the crests take the form of high flanges
The latter (up to 6 /z. in height) are membranous, thin and always perforate.
The number of apical plates always appears to be six. Plate I', equivalent to
the first apical plate, is elongate and abuts against the anterior end of the sulcus.
The crest arrangement on the ventral surface appears to be practically constant and
is characteristic of this species. The crests limiting the plates F, I", II", III", IV",
VII" and VIII" are always constant in position. Plates F" and IF" are reduced
due to the presence of a posterior intercalary plate. Plate IF" always possesses a
crest passing diagonally across it and the crest between plates IIF" and IV" is of a
A B
FIG. ii. Cribroperidinium intricatum sp. nov. A. Ventral surface of Holotype,
(X 700). B. Reconstruction to show Gonyaulax-type tabulation (x 700).
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i 127
characteristic right angle shape. The positions of the crests on the dorsal surface
of the epitract are fairly constant, only varying in minor details. The large plate
V" is usually detached in archaeopyle formation and possesses a semi-circular crest
from which radiate a small number of other crests (PI. 2, fig. 3). These pass over
the boundary of plate V" to continue on adjacent parts of the epitract. The posi-
tions of the crests on the dorsal surface of the hypotract appear to be less constant
in position. Their predominent direction is parallel to the longitudinal axis of the
shell; sometimes a series of posterior circle plates may be present. Antapical
plates, if present at all, are very reduced and obscured by the crestal membranes.
The cingulum is narrow (c. 6/z in width), slightly laevo-rotatory, and tends to
possess a thicker wall than is usual for the remainder of the shell. The sulcus only
projects onto the epitract for a short distance, being considerably larger and wider
on the hypotract. It is always divided into a number of posterior ventral plates by
reduced crests.
REMARKS. Figs nB, I2B have been prepared from the holotype and one
other specimen, omitting the additional crests, to show the basic Gonyaulax-type
tabulation.
C. intricatum may be differentiated from C. orthoceras (Eisenack), C. muderongensis
(Cookson & Eisenack), Gonyaulacysta apionis and G. diaphanis by its more spherical
form, details of crest arrangement and the absence of spines. C. edwardsi (Cookson
& Eisenack) is most similar, being almost spherical, but possesses a very well
developed, stiff apical horn and the crests on the ventral surface are arranged differ-
ently. In particular the diagonal crest on plate II'" is absent. Gonyaulax sp. (Gorka
1963) from the Cenomanian of Poland is very similar and may be conspecific
with C. intricatum.
A B
FIG. 12. Cribroperidinium intricatum sp. nov., A. Dorsal surface of Holotype with
archaeopyle (X 700). B. Reconstruction to show Gonyaulax-type tabulation (X 700).
128 CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
OCCURRENCE. C. intricatum is very rare in the British Cenomanian and has been
observed only in samples FM 840, CB I and CB 7. At Escalles it is infrequent in
sample E 201, very rare in sample E 195 and lacking elsewhere. In the Saskatche-
wan sample Sas 1023 it is common, in sample Sas 1084 rare and is lacking in all the
other North American samples. It is interesting to record that this species was
observed in the Albian sample FM 886 from Fetcham Mill. Thus C. intricatum has
a range from Albian to Lower Cenomanian in the examined material.
OTHER SPECIES
The following species are here attributed to the genus Cribroperidinium on the
basis of overal shape, and number and position of periphragm crests: —
Cribroperidinium orthoceras (Eisenack 1958) comb, nov., 1958 Gonyaulax orthoceras Eisenack,
Neues. Jb. Geol. Palaont., Abh., 106 (3) 388: pis. 21, figs. 3-14; 24, fig. i; text-figs. 2, 3.
Cribroperidinium edwardsi (Cookson & Eisenack 1958) comb, nov., 1958 Gonyaulax edwardsi
Cookson & Eisenack: Proc. R. Soc. Viet., 70 (i), 32-33; PL III, figs. 5, 6, text-fig. 7.
Cribroperidinium muderongensis (Cookson & Eisenack 1958) comb, nov., 1958 Gonyaulax
muderongensis Cookson & Eisenack: Proc. R. Soc. Viet., 70 (i), 32; PI. Ill, figs. 3, 4, text-fig. 15.
Gonyaulacysta apionis (Cookson & Eisenack 1958) and G. diaphanis (Cookson &
Eisenack 1958), both from the Lower Cretaceous of Australia, are of similar appear-
ance to forms included in Cribroperidinium and may at a later date be transferred
to this genus.
Eisenack (1958 text-figs. 2, 3) figured the ventral surface tabulation of C. ortho-
ceras omitting, or dotting in, some of the crests which are seen to be present on the
photographs of the same specimens. Thus a true representation of the crestal
positions was not given, somewhat misleading later workers. These figures have
been redrawn (Figs. I3A, B) from the photographs and show the remarkable similarity
between the ventral surface of C. orthoceras and that of C. intricatum. For this
reason the former species has been transferred to Cribroperidinium and the diagnosis
emended. Eisenack does in fact compare and contrast his species with the
reattributed Australian forms above, also with Gonyaulactysa wetzeli (Lejeune
Carpentier 1939) and G. obscura (Lejeune-Carpentier 1946), all of which he con-
siders to be in the same group. The latter two species, however, have a typical
Gonyaulacysta tabulation and must remain in that genus.
Cribroperidinium orthoceras (Eisenack) emend.
(Figs. I3A, B)
1958 Gonyaulax orthoceras Eisenack : 388, pi. 21, figs. 3-14, pi. 24, fig. i ; text-figs. 2, 3.
!959 Gonyaulax orthoceras Eisenack; Gocht : 54, pi. 5, figs. 12, 13.
1961 Gonyaulax orthoceras Eisenack; Alberti : 6, pi. n, figs. 1-3.
1963 Gonyaulax orthoceras Eisenack; G6rka : 30, pi. 3, figs. 1-4.
1965 Gonyaulax orthoceras Eisenack; Baltes : 12, pi. 3, figs. 95-99.
EMENDED DIAGNOSIS. Shell ovoidal, moderately thick-walled, bearing strong,
thorn-like apical horn constituting approximately one quarter of shell length. Shell
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 129
wall granular bearing tubercles. Distinct tabulation marked by strong, low crests
delimiting large number of plates. Plate IF" crossed diagonally by low crest
and operculum possessing semi-circular crest. Cingulum narrow, devoid of plate
boundaries.
HOLOTYPE. The specimen illustrated by Eisenack (1958, pi. 21, fig. 5) from Pre-
paration Ob. Apt. No. 32. Aptian glauconitic limestone, Deutschen Erdol A. G.,
Erdolwerke Holstein boring Marne, Feld Heide, North Germany, at 7617 metres
depth.
REMARKS. The emended diagnosis excludes those forms described by Sarjeant
(1966) from the Speeton Clay which are to be described elsewhere under a new specific
name.
Genus CARPODINIUM Cookson & Eisenack, 1962
Carpodinium obliquicostatum Cookson & Hughes
(PL i, figs. 3, 4)
1964 Carpodinium obliquicostatum Cookson & Hughes : 48, pi. 6, figs. 1-6.
1967 Carpodinium obliquicostatum Cookson & Hughes; Clarke & Verdier: 23, pi. 2, figs. 4, 5.
DESCRIPTION. The shell is elongate-ovoidal and bears relatively high sutural
crests. The latter are smooth or slightly granular and the distal margin may be
entire or spinous. The short apical horn is a prolongation of one of the larger crests
and is typically asymmetrically placed. The periphragm of the shell possesses an
unusual ornamentation — small areas of triangular to polygonal shape, composed
of thick periphragm, are separated by narrow anastomosing " canals " where the
periphragm is unthickened or perhaps absent (PL r, fig. 4).
The cingulum is marked by indentations in some of the crests and is not apparent
on the shell surface. The sulcus, delimited by crests, is occasionally observable
and extends the length of the shell, being narrow near the apex and widening pos-
teriorly. The precingular and postcingular plates, probably six in each series, are
very elongate and difficult to discern because of the obscuring nature of the high
crests. A precingular archaeopyle is commonly present. Four apical plates and a
single antapical plate appear to be present. Intercalary plates were not observed.
DIMENSIONS. Range of observed specimens: overall length 56(69-5) 82 //,, overall
width 33(43-8) 57 /*, height of crests 6-15 ^u,. Number of specimens measured, 16.
REMARKS. The Cenomanian specimens studied resemble the type material from
the Upper Albian and Lower Cenomanian of Cambridgeshire in all respects.
OCCURRENCE. C. obliquicostatum is a rare to very rare species found at all
horizons throughout the Cenomanian of Fetcham Mill and Compton Bay and in
three samples from Escalles (E 195, E 189 and E 159).
Genus ELLIPSODINIUM Clarke & Verdier 1967
REMARKS. A number of microplankton genera have been described as possessing
a reticulate shell wall, occasionally with an outer membrane but only rarely with any
130 CENOMANIAN NON-CALCAREOUS MI CROPL AN KTON, i
signs of tabulation. In all the described forms possessing a cingulum the archae-
opyle is apical. Hence the combination of numerous crests, a cingulum and a
precingular archaeopyle differentiates Ellipsodinium from all previously described
genera.
Ellipsodinium rugulosum Clarke & Verdier
(PI. 3. fig. i ; Figs. 140, D)
1967 Ellipsodinium rugulosum Clarke & Verdier: 69, pi. 14, figs. 4-6, text-fig. 29.
DIMENSIONS. Range of observed specimens : shell length 30 (37 . 7) 46 ju., shell width
25 (33-6) 40 p, maximum height of crests 1-5 (2-4) 3-5 ju,. Number of specimens
measured, 20.
DESCRIPTION. The crests are thin, occasionally perforate lamellar structures which
thicken slightly before joining the shell surface. The cingulum may be delimited by
a pair of crests, or in the absence of cingular crests, crests may terminate abruptly
at its borders. Rarely crests traverse the cingulum. The sulcus is not obvious
because of the nature of the elongate areas outlined by the crests. Apical and
antapical processes or horns are absent. The precingular archaeopyle is sub-
triangular in outline.
OCCURRENCE. E. rugulosum is a rare to fairly common species at all horizons
throughout the Cenomanian of Fetcham Mill, Compton Bay and Escalles. It was
not observed in the North American material.
Cyst-Family PAREODINIACEAE Gocht emend. Sarjeant & Downie 1966
Genus APTEODINIUM Eisenack 1958
REMARKS. Members of this genus have occasionally been observed in the Euro-
pean Cenomanian and have been placed in A . granulatum. Rarely plate boundaries
may be discernable. This fact together with the overall shape and the well developed
precingular archaeopyle indicate that this genus is closely related to Gonyaulacysta
and at a future date may be transferred to the Cyst-Family Gonyaulacystaceae.
Apteodinium granulatum Eisenack
(PI. 3, figs. 5, 6)
?I935 Palaeoperidinium ventriosum O. Wetzel; Deflandre : 228, pi. 5, fig. 5; pi. 6, figs. 9, 10.
?i936& Palaeoperidinium ventriosum O. Wetzel; Deflandre : 27, pi. 5, figs. 1-4.
?i936a Palaeoperidinium ventriosum O. Wetzel; Deflandre: fig. 100.
1958 Apteodinium granulatum Eisenack : 386, pi. 23, figs. 8-14, text-fig, i.
1958 Apteodinium granulatum Eisenack; Gocht : 64, pi. 5, fig. 2.
1961 Apteodinium granulatum Eisenack; Alberti : 24, pi. 4, figs. 4-6.
1963 Apteodinium granulatum Eisenack; Baltes. : 584, pi. 4, fig. u.
DESCRIPTION. Shell subspherical with a moderately thick wall (1-2-5 //,)
possessing a stout conical horn. The wall is densely granular, the granules often
being elongated into fine, short processes which are linked laterally thus covering
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 131
the shell surface with an intricate network of a matted furry appearance. The
cingular boundaries are always visible as slight thickenings of the periphragm.
Very rarely similar but more reduced thickenings indicate the presence of plate
boundaries. A precingular archaeopyle is commonly developed.
DIMENSIONS. Range of observed specimens: overall length 42 (48-2) 53 /z, width
31 (38-6) 44 p., length of apical horn 5 (6-1) 8 /*. Number of specimens measured, 9.
REMARKS. The Cenomanian specimens are very similar in appearance to the
type material described by Eisenack from the Aptian of Germany. The matted,
furry appearance was not described but when this feature is only slightly developed
the shell wall merely appears to be very granular. The Aptian and Cenomanian
examples of A . granulatum appear to be comparable to Palaeoperidinium ventriosum
O. Wetzel as illustrated by Deflandre (1935, 1936^, b). The holotype of this species
has been re-examined by Lejeune-Carpentier (1946) and a distinct tabulation des-
cribed. Deflandres' forms which do not possess a tabulation are hence here ten-
tatively reattributed to A . granulatum.
OCCURRENCE. Only two specimens have been recorded from the English Ceno-
manian, both from sample FM 690. At Escalles two specimens have been recorded
from both samples E 183 and E 153, and three from sample E 165. A. granulatum
is fairly common in sample FM 886 (Albian) from Fetcham Mill but is absent in the
North American material and from sample FM 520 (Turanian).
Genus TRICHODINIUM Eisenack & Cookson emend. Clarke & Verdier
REMARKS. This genus differs from Exochosphaeridium Davey, Downie, Sarjeant
& Williams (1966) by the presence of a well developed cingulum and by the shorter
spines.
Trichodinium castaneum (Deflandre)
PI. n, figs. 1-3
J935 Palaeoperidinium castanea Deflandre : 49, pi. 6, fig. 8.
19366 Palaeoperidinium castanea Deflandre; Deflandre : 25, pi. 16, figs. 1-4.
19360 Palaeoperidinium castanea Deflandre; Deflandre: fig. 99.
19526 Palaeoperidinium castanea Deflandre; Deflandre: fig. 96.
19626 Palaeoperidinium castanea Deflandre; Cookson & Eisenack : 489, pi. 3, figs. 9-11.
1964 Palaeoperidinium castanea Deflandre; Cookson & Hughes : 49, pi. 5, fig. 14.
1967 Trichodium castanea (Deflandre) Clarke & Verdier: 19, pi. i, figs, i, 2.
DESCRIPTION. The shell is subspherical with occasionally a small apical horn or
a tuft of apical spines. The shell wall is slightly punctate and bears numerous small
spines. These are solid, often bifurcate either proximally or distally, and typically
terminate in a small bifurcation. Rarely they may be acuminate. The cingulum
(3-5 fj, in width) is marked by two parallel lines of thickening along which spines are
concentrated. Lines of similar thickening sometimes occur perpendicularly to the
cingulum and are probably sutural. A sulcus has not been observed. A precingular
archaeopyle is commonly present.
132 CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i
DIMENSIONS. Range of observed specimens: shell diameter 35 (49-0) 64 /n, length
of spines I (3-3) 5 /x. Number of specimens measured, 16.
REMARKS. The Cenomanian specimens resemble the type material from the
Upper Cretaceous of France in all respects. The presence of a precingular archaeo-
pyle, an apical prominence or apical spines, and a well developed cingulum indicate
that this species should be placed in Trichodinium. T. intermedium Eisenack &
Cookson, from the Aptian to Lower Albian of Australia, is very similar but is larger
(shell diameter 69-90 /A).
OCCURRENCE. T. castaneum is a rare species occurring at most horizons throughout
the Cenomanian of Fetcham Mill, Compton Bay and Escalles. It has not been
recorded from the North American samples. This species has a recorded strati-
graphic range from the Aptian to the Senonian (Deflandre).
Cyst-Family MIGRODINIACEAE Eisenack emend. Sarjeant & Downie 1966
Genus MICRODINIUM Cookson & Eisenack emend. Sarjeant 1966
REMARKS. Microdinium differs from Gonyaulacysta primarily in possessing a
single apical plate which is detached in archaeopyle formation. Gorka (1965), in des-
cribing Upper Jurassic assemblages, erected the genus Tetrasphaera which is diagnosed
as having a feebly marked tabulation and short spines on the plate boundaries. Gorka
has since stated (personal communication with Dr. W. A. S. Sarjeant) that this
genus differs from Microdinium only in these two respects. However, it follows
that if the tabulation is not clear it is difficult to compare this new genus with
Microdinium or any other genus possessing a tabulate cyst, since the diagnosis of
these genera is primarily based on the observed tabulation. Also, spines have been
observed on the plate boundaries of Microdinium, even in the type species. Thus
it would appear that Tetrasphaera at the moment is not precisely defined.
Microdinium cf. ornatum Cookson & Eisenack.
(PI. 4, fig. 5; Figs. I3C, F)
ig66fl Microdinium cf. ornatum Cookson & Eisenack; Sarjeant : 149, pi. 16, figs. 3-6, text-fig. 38.
1967 Microdinium ornatum Cookson & Eisenack; Clarke & Verdier: 66, pi. 5, figs. 11-14.
DESCRIPTION. The shell is subspherical to ovoidal possessing a smooth body wall
ornamented by a few large tubercles. The latter may be flat or distinctly concave
distally. In two specimens small tubercles were seen to delimit a plate in the
posterior portion of the ventral area. The plates are bordered by short, broad
projections (Fig. I3F), which may be isolated or united distally.
DIMENSIONS. Range of observed specimens: shell length 31-34 //,, width 23-31 /z,
height of crests 1-2 /*. Number of specimens measured, 6.
REMARKS. M . cf. ornatum differs from M. ornatum Cookson & Eisenack (19600)
in that (i) the small cingular plate, ventral to plate 6c, is absent or has a very
reduced ventral suture, and (ii) the plates are not bordered by ledges, which are
sometimes perforate, but by isolated tubercles and spines.
CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i 133
OCCURRENCE. M. cf. ornatum is a rare species in the material examined being
found only in the following samples: FM 810, FM 670, FM 650, FM 520 (Turonian),
E 189 and E 153. In Australia M. ornatum has a stratigraphic range from the Albian
to the Lower Turonian.
Microdinium setosum Sarjeant.
(PI. 2, fig. 4; Fig. i3H)
1966 Microdinium setosum. Sarjeant : 151, pi. 16, figs. 9, 10; text-fig. 39.
1967 Microdinium echinatum Clarke & Verdier: 64, pi. i, figs. 9, 10, text-fig. 26.
DESCRIPTION. The shell is spherical to ovoidal and possesses a well developed
tabulation. The shell surface is either lightly or coarsely granular. The two plates
observed by Sarjeant in the medial region of the sulcus are not always delimited.
The sutural crests are often relatively high and give rise to numerous, well developed
thorn-like spines (Fig. I3H).
DIMENSIONS. Range of type material: shell length 25 (29-3) 37 p., width 21 (26-2)
31 p, maximum height of crests 1-5 (3-8) 7 /x. Number of specimens measured, 22.
REMARKS. In general form M. setosum is similar to M. ornatum but differs in the
presence of spiny crests, a dense granulation and, slightly, in the tabulation exhibited.
OCCURRENCE. M. setosum is an infrequent to fairly common species at most
horizons throughout the Cenomanian of Fetcham Mill, Compton Bay and Escalles.
It has also been recorded from the Albian of Fetcham Mill.
Microdinium distinctum sp. nov.
(PI. 2, figs. 9-11 ; Figs. 130, E. I)
1967 Microdinium ornatum Cookson & Eisenack; Clarke & Verdier: pi. 5, figs. 11-12.
DERIVATION OF NAME. Latin, distinctus, different — with reference to the
distinctive appearance of this species.
DIAGNOSIS. Shell subspherical, thick-walled, smooth. Sutural crests well
developed bearing short, broad, flat-topped spines. Reflected tabulation i, oa, 6"
6"', ip, i""; plates i" and 6" very reduced and cingular plates absent. Cingulum
broad, weakly laevo-rotatory.
HOLOTYPE. G.S.M., slide PF 3989, (i). Lower Chalk, H.M. Geological Survey
Borehole, Fetcham Mill, Surrey at 730 ft. depth. Upper Cretaceous (Cenomanian).
DIMENSIONS. Holotype: shell length 36^, width 37 p, height of crests c. 2,\i.
Range: shell length 29-36^, width 30-37 //., height of crests 2-2-5^. Number of
specimens measured, 7.
DESCRIPTION. The shell wall is relatively thick, c. i /z, and develops pronounced
ridges at the plate boundaries. The sutural spines are closely set, very broad and
flat distally.
134
GENOMANIAN NON-CALCAREOUS MICROPLANKTON, i
H
FIG. 13. Cribroperidinium orthoceras (Eisenack). A. Specimen illustrated by Eisenack
(1958, text-fig. 3) redrawn to show full crestal arrangement. B. Holotype (Eisenack
1958, text-fig. 2) redrawn to show full crestal arrangement. Microdinium cf. ornatum
Cookson & Eisenack, C. Ventral surface of Holotype (X 700). Microdinium distinctum
sp. nov., D. Ventral surface of Holotype (x 700), E. Dorsal surface of Holotype (X 700).
Microdinium cf. ornatum Cookson & Eisenack, F. Sutural spines. Microdinium vario-
spinum sp. nov., G. Sutural spines. Microdinium setosum Sarjeant, H. Sutural spines.
Microdinium distinctum sp. nov., I. Sutural spines.
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 135
The precingular plates are generally smaller than the postcingular plates. Plates
i" and 6" are small and appear as slight projections in the ventral area. This is
widest posteriorly and is open anteriorly. The apical plate is six-sided and is lost
in archaeopyle formation. The archaeopyle possesses slits extending posteriorly
between the precingular plates.
REMARKS. M . distinctum may be distinguished from all other species of Micro-
dinium by the thick, smooth shell wall, the form of the precingular plates and the
absence of cingular plates. It is considered that the absence of the latter does not,
at present, warrant the erection of a new genus.
OCCURRENCE. This species is very rare, being recorded only seven times, from the
following samples: FM 790, FM 730, FM 710, FM 690, £153 and €617. It has
never been observed in the lower horizons of the Cenomanian.
Microdinium variospinum sp. nov.
(PI. 2, figs. 5, 6; Fig. i3G)
DERIVATION OF NAME. Latin, varius, different; spinosus, spine — with reference
to the variable appearance of the spines.
DIAGNOSIS. Shell subspherical to ovoidal; shell wall thin, smooth, granular or
lightly reticulate. Sutural crests low, bearing small number of variably shaped
spines. These may be simple tubercles to complex bifurcating protrusions. Re-
flected tabulation i', oa, 6", (6c), 6'", rp, i""; cingular plate boundaries very faint
or absent. Cingulum weakly laevo-rotatory.
HOLOTYPE. B.M. (N.H.) V. 51981 (i). Lower Chalk, Bureau de Recherches
Geologiques et Minieres Borehole, Escalles, Pas de Calais at 165 metres depth.
Upper Cretaceous (Cenomanian) .
DIMENSIONS. Holotype: shell length 24^1, width 23/4, length of spines 1-1*5/4
Range: shell length 20 (23-6) 27/4, width 17 (20-0) 23/i, length of spines 1-3 /u,.
Number of specimens measured, 9.
DESCRIPTION. The cingular plate boundaries are usually absent, however, one
Saskatchewan specimen had them lightly defined. The sutural spines are few in
number and tend to be concentrated at the posterior of the shell (PI. 2, fig. 5). They
are stout and may be either pointed or flattened distally (Fig. 130).
REMARKS. The characteristic features of M. variospinum are the nature of the
shell wall, the form of the spines and the lack of well defined cingular plate bound-
aries. These characters together differentiate this species from all other described
species.
OCCURRENCE. M. variospinum is a rare species found at all horizons, save the
lower three, at Escalles, and in sample CB i, CB 7 and Sas 1084. It has not been
recorded from Fetcham Mill.
GEOL. 17, 3 IO
136 CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
Microdiniutn veligerum (Deflandre) comb. nov.
(PI. 3, fig- 4i PL 4, fig- 4)
1937 Micrhystridium veligerum Deflandre : 81, pi. 12, fig. 9.
1943 Ceratocorys veligera (Deflandre) Lejeune-Carpentier : 22, text-figs. 1-6.
19520 Ceratocorys veligera (Deflandre) Deflandre : 120, text-fig. 102.
i952b Ceratocorys veligera (Deflandre) Deflandre: text-figs. 304 A-C.
1967 Eisenackia crassitabulata Deflandre & Cookson; Clarke & Verdier: 64, pi. 8, figs. 4-6.
DESCRIPTION. The shell is ovoidal, densely granular and bears a number of high
crests delimiting a tabulation. The crests are distinctive, being i to 5 /u in height,
with typically a smooth outer margin. They consist of two membranes, joined
distally and diverging proximally to form a broad base, I to 3 p, wide to the crest.
There is between the two membranes a crestal cavity which is occasionally sub-
divided by septa, particularly where two crests diverge. Here a conical chamber is
usually found. The reflected tabulation appears to be i', ?5", 6c, 6'", ip, i"".
The hypotract is considerably larger than the epitract, the latter being devoid of
crests and usually possessing a pentagonal apical archaeopyle. The shape of the
latter is the only indication that there are five precingular plates. The cingulum is
broad and does not appear to be spiral. Plate i'", and to a lesser extent 2'", are
reduced to accommodate the posterior intercalary plate. The remaining four
postcingular plates are large and there is a single large antapical plate. The sulcus
is very narrow just posterior to the cingulum and then widens rapidly towards the
antapex. The sulcus extends onto the epitract where it sometimes bears five small
sulcal plates.
DIMENSIONS. Range of observed specimens: shell length 28 (31-5) 38 /*, width
25 (28-2) 32 //,. Number of specimens measured, 13.
REMARKS. Lejeune-Carpentier (1943) placed this species, originally observed in
the Upper Cretaceous of France, in the genus Ceratocorys Stein (1883) on the basis
of its similarity to motile dinoflagellates contained in this genus. However, it is a
cyst possessing an apical archaeopyle and should not be attributed to a motile
dinoflagellate genus. Thus this species is here transferred to Microdinium on the
basis of the tabulation, apical archaeopyle, reduced size of epitract compared with
the hypotract and overall small size. M. veligerum does, however, differ slightly
from the other species in this genus by the apparent absence of crests on the epitract
and the probable presence of five precingular plates instead of six.
Eisenackia crassitabulata as illustrated by Clarke & Verdier (1967) is undoubtedly
M. veligerum. The former, as originally described from the Australian Lower
Tertiary, is of different overall form and is considerably larger (72-78 by 55-67 /*) .
The size of the specimen illustrated by Clarke & Verdier is approximately 30 by 32 /£
and is thus comparable to the Cenomanian specimens of M. veligerum. E. crassi-
tabulata has been recorded from the Lower Tertiary and also from the Maestrichtian
of South Africa by the present author. M. irregulare Clarke & Verdier (1967)
appears to be very similar to M. veligerum and any definite distinction is not apparent.
OCCURRENCE. The Cenomanian forms examined resemble the specimens illus-
trated by Lejeune-Carpentier in all respects. M. veligerum is an infrequent to
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i 137
common species at all horizons, save two, at Fetcham Mill, Compton Bay and
Escalles. These two horizons are basal Cenomanian, FM 840 and CB i, and it was
not recorded by Cookson & Hughes (1964) from the Upper Albian/basal Cenomanian
of Cambridgeshire. It is also absent in the North American material and has not been
described from Australia. This species is present in sample FM 520 of Turonian
age. M. veligerum thus appears for the first time just above the base of the Ceno-
manian and extends into the Turonian but is, apparently, of restricted geographical
distribution.
fMicrodinium crinitum sp. nov.
(PL 2, figs. 7, 8)
1967 Cometodinium obscurum Deflandre & Courteville; Clarke & Verdier: pi. 10, fig. 3: pi. n,
fig. 9-
DERIVATION OF NAME. Latin, crinitus, hairy — with reference to the numerous
hair-like spines.
DIAGNOSIS. Shell subspherical, periphragm granular and giving rise to numerous,
fine, flexuous spines. Sutural crests low, bearing numerous spines. Cingulum
wide, composed of elongate plates. Epitract smaller than hypotract. Archaeopyle
not normally visible.
HOLOTYPE. G.S.M. slide PF 3990(1). Lower Chalk, H.M. Geological Survey
Borehole, Fetcham Mill, Surrey at 690 feet depth. Upper Cretaceous (Cenomanian.)
DIMENSIONS. Holotype: shell diameter 27 by 28 p, length of spines c. 12 /JL.
Range: shell diameter 24 (30-1) 38/1,, length of spines 6-19 /x. Number of specimens
measured, 18.
DESCRIPTION. The periphragm granules, which are evenly spaced on the shell
surface, are up to 0-5 /j, high, and often form the bases of the hair-like spines. The
latter tend to be especially concentrated along the sutures, and because of this
tendency and the spherical form of the shell, it has not been possible to fully for-
mulate a tabulation. However, precingular, cingular and postcingular plates are
quite obvious when the orientation is favourable, the precingular plates being smaller
than the postcingular plates. The cingulum is broad, c. 5 /x. The archaeopyle,
although it has not been observed, is probably apical.
REMARKS. The numerous hair-like spines and the tabulation easily distinguish
1M. crinitum sp. nov. from all previously described forms of dinoflagellate cysts.
The overall shape, the small size and the fact that the epitract is smaller than the
hypotract all indicate that this species is closely related to the genus Microdinium.
However, plate spines have not been recorded in Microdinium, although a granula-
tion has, and since the tabulation has not been elucidated in the present species it is
only placed tentatively in this genus.
OCCURRENCE. 1M. crinitum is infrequent at all horizons throughout the Ceno-
manian of Fetcham Mill, Compton Bay and Escalles and is also present in the Albian
sample from Fetcham Mill, sample FM 886. It has only once been recorded at
Saskatchewan, in sample Sas 835 (Cenomanian).
138 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
Genus HISTIOCYSTA nov.
DERIVATION OF NAME. Greek, histos, mesh or network; kystis, sac or cell — with
reference to the reticulate ornamentation on the shell surface.
DIAGNOSIS. Proximate cysts, spherical to subspherical ; shell wall composed of
two layers; outer layer giving rise to low crests. Crests reflecting Gonyaulax—
type tabulation and coarse, subsidiary reticulation within plate boundaries. Sutural
crests better denned than crests of subsidiary reticulation. Archaeopyle apical with
angular margin. Operculum probably single apical plate.
TYPE SPECIES. Histiocysta palla sp. nov. Lower Chalk (Cenomanian) ; England.
REMARKS. The reasonably well denned tabulation, the plate ornamentation and
the apical archaeopyle easily distinguish Histiocysta from all previously described
genera. The most similar genera are Ellipsoidictyum Klement (19606) and Dic-
tyopyxidia Eisenack (1961), both from the Upper Jurassic. Both genera possess
an apical archaeopyle and a cingulum but a distinct Gonyaulax-type tabulation is
absent. However, it seems probable that Histiocysta is genetically related to these
two genera. Although the precise tabulation of Histiocysta cannot be elucidated
the presence of precingular, cingular and postcingular series of plates together with
an apical archaeopyle indicate that this genus belongs to the Family Microdiniaceae.
Histiocysta palla sp. nov.
(PI. i, figs. 5, 6; Figs. I4A, B)
1939 Micrhystridium sp.? Deflandre & Courteville : pi. 3, fig. 4.
DERIVATION OF NAME. Greek, palla, ball — with reference to the more or less
spherical shape of this species.
DIAGNOSIS. Shell spherical to subspherical, thin-walled, periphragm smooth and
forming a reticulate network of crests. Network consisting of reflected dinoflagel-
late tabulation with central region of each plate occupied by coarse but simple
reticulation.
HOLOTYPE. G.S.M. slide PF 3052 (2). Lower Chalk, H.M. Geological Survey
Borehole, Fetcham Mill, Surrey at 650 feet depth. Upper Cretaceous (Cenomanian) .
PARATYPE. G.S.M. slide PF 3991(1). Lower Chalk, H.M. Geological Survey
Borehole, Fetcham Mill, Surrey at 710 feet depth. Upper Cretaceous (Cenomanian).
DIMENSIONS. Holotype : shell diameter 30 by 33 p, height of crests 4-5 /*. Para-
type: shell diameter 26 by 29^, height of crests c. 2-5 /A. Range: shell diameter
25 (31-8) 38 /z, height of crests 1-5 /z. Number of specimens measured, 19.
DESCRIPTION. The precingular and postcingular plates may be observed on most
specimens but the exact tabulation has not, as yet, been elucidated. The central
region of each plate is occupied by a coarse, subpolygonal reticulation which occasion-
ally extends to the plate boundaries (Figs. I4A, B). The cingular region is clearly
defined by crests and encircles the shell. Cingular plates are not usually discernible,
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
139
the entire region being occupied by a coarse reticulation. An apical archaeopyle
with an angular margin is typically present, the six-sided operculum often remaining
attached to the shell.
REMARKS. The combination of apical archaeopyle, tabulation and reticulation
makes H. pdlla an easily recognizable species distinct from all previously described
B
C D
FIG. 14. Histiocysta palla sp. nov., A. Lateral view of Holotype showing attached oper-
culum, apical archaeopyle, precingular plates and cingulum (X 13°°)- B. Lateral
view of Holotype showing well defined plate boundaries (X 13°°)- Ellipsodinium
rugulosum Clarke & Verdier, C. Lateral view showing partially detached operculum and
cingulum (X 1300). D. Lateral view (X 1300).
i4o CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
forms. This species was figured, but not described, by Deflandre & Courteville
(1939) as Micrhystridium sp.? from the Senonian. Membranilarnax cf. pterosper-
moides Deflandre (1937) is of similar form but does not possess a reticulation.
OCCURRENCE. H. palla is found throughout the European Cenomanian, except
for the lowermost horizons. It is fairly common at Fetcham Mill but rare at Comp-
ton Bay and Escalles. It has not been observed in the North American samples.
Cyst-Family FROMEACEAE Sarjeant & Downie 1966
Genus FRO ME A Cookson & Eisenack 1958
REMARKS. A number of specimens referable to the type species, F. amphora, do
not appear to possess a cingulum. The absence of a cingulum makes this genus
similar to Chytroeisphaeridia Sarjeant (1962). They differ, however, in that the
archaeopyle of Fromea has a rounded margin, and in the elongate shape typical of
the latter genus.
Fromea amphora Cookson & Eisenack
(PI. 3, figs. 2, 3)
1958 Fromea amphora Cookson & Eisenack : 56, pi. 5, figs. 10, u.
19666 Fromea amphora Cookson & Eisenack; Sarjeant : 209, pi. 22, fig. 4; pi. 23, fig. 3 (see also
for earlier references).
DESCRIPTION. The shell is ovoidal, thick-walled (2-3 p,) and typically possesses
an apical archaeopyle with a rounded margin. In one specimen (PL 3, fig. 3) the
apical region is still attached and may be seen to be perfectly rounded. A cingulum
was not observed in any of the specimens.
DIMENSIONS. Range of observed specimens: shell length 56 (72-5) 85^, width
47 (60) 71 /A. Number of specimens measured, 6.
REMARKS. The Cenomanian specimens are identical with the type material from
the Aptian-Cenomanian of Australia except that the cingulum is absent. Cookson
& Eisenack (1958) state, however, that the cingulum may be rather faint and it is
probable that the European forms fall within the range of variation for this species.
F. amphora has been recorded from the Barremian of England by Sarjeant (19666).
The specimens described by Sarjeant, like the Cenomanian forms, do not possess a
cingulum. Maliavkina et al. (1961) describes some very similar, but rather smaller,
forms from the Maestrichtian of Siberia, calling them Chrysomonadinael . These
forms possess the typical rounded archaeopyle of this genus.
OCCURRENCE. Five specimens have been recorded from Fetcham Mill, from
samples FM 810, 770, 750 and 650, and one specimen from Escalles, sample E 177.
Genus CHYTROEISPHAERIDIA Sarjeant 1962
REMARKS. Chytroeisphaeridia and Canningia Cookson & Eisenack (19606) are
similar and probably fairly closely related. The latter is usually more polygonal,
has an apical horn, and sometimes the vestiges of a cingulum.
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 141
Chytroeisphaeridia euteiches sp. nov.
(PL 3, figs. 8, 9)
DERIVATION OF NAME. Greek, euteiches, well- walled — with reference to the stout
wall of this species.
DIAGNOSIS. Shell subspherical; shell wall thick and densely granular. Angular
apical archaeopyle typically present.
HOLOTYPE. B.M. (N.H.) V. 51982 (2). Lower Chalk, Bureau de Recherches Geo-
logiques et Minieres Borehole, Escalles, Pas de Calais, at 159 metres depth. Upper
Cretaceous (Cenomanian).
DIMENSIONS. Holotype: shell length 53^, width 59 /*. Range: shell length
48-60 jit, width 49-59 p. Number of specimens measured, 6.
DESCRIPTION. The shell wall is thick (2-3 /*) and may be composed of two layers.
If the wall is bipartite then the inner layer is thin, the outer layer making up almost
the entire wall thickness. This layer appears to be composed of minute cellular
elements and is densely granular on the surface. The apical archaeopyle, when
developed, is angular with small slits passing posteriorly from its margin between each
precingular plate. A sulcal notch is also present (PI. 3, fig. 8). Plate boundaries
and cingulum are not discernible.
REMARKS. This species is a simple, subspherical shell possessing an apical
archaeopyle. One species, C. chytroeides Sarjeant (1962), from the Upper Jurassic
of England, differs from C. euteiches in having a fairly thin and only slightly granular
shell wall. Chytroeisphaeridia sp. Sarjeant (19656) is similar in size and also granular
but does not possess a thick shell wall. Canningia rotundata Cookson & Eisenack
(1961) is also similar but tends to have a polygonal shell with a small apical horn.
OCCURRENCE. C. euteiches has been recorded from two horizons only, E 165, & E
159 from Escalles, where it is infrequent.
Genus CASSICULOSPHAERIDIA nov.
DERIVATION OF NAME. Latin, cassiculus, hunting-net; sphaera, ball — with
reference to the surface reticulation of the shell.
DIAGNOSIS. Proximate cysts; shell spherical to subspherical, composed of two
layers, without apical or antapical protuberances. Periphragm giving rise to low
crests or membranes which form a reticulate pattern. Tabulation absent. Archae-
opyle apical with angular margin.
TYPE Species. Cassiculosphaeridia reticulata sp. nov. Lower Chalk (Ceno-
manian); France.
REMARKS. The surface reticulation of this genus is very similar to that of
Ellipsoidictyum cinctum Klement (1960). Both possess an apical archaeopyle, but
whereas in Cassiculosphaeridia all signs of a tabulation are absent, in Ellipsoidictyum
there is an obvious cingulum. Dictyopyxidia Eisenack (1961) is also very similar
but possesses a cingulum and sulcus.
142 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
Cassiculosphaeridia reticulata sp. nov.
(PL 3, fig- 7: PL 4, %• 3)
DERIVATION OF NAME. Latin, reticulatus, net-like — with reference to the
reticulate pattern formed by the periphragm crests.
DIAGNOSIS. Shell spherical to subspherical. Shell surface bearing low ridges,
forming a coarse reticulation, from which arise fine membranous crests. Shell wall
lightly to densely granular.
HOLOTYPE. B.M. (N.H.) ¥.51981 (4). Lower Chalk, Bureau de Recherches
Geologiques et Minieres Borehole, Escalles, Pas de Calais, at 165 metres depth.
Upper Cretaceous (Cenomanian) .
DIMENSIONS. Holotype: shell diameter 38 by 38/11, height of crests 4 to 6 p.
Range : shell diameter 33 (43-6) 55 p,, maximum height of crests 3 (7-3) II /n. Number
of specimens measured, 15.
DESCRIPTION. The areas delimited by the low ridges are typically subpolygonal
but may be of irregular shape. They vary considerably in size, from 2 to io/u, in
diameter. The membranous crests are very fine and tend to be flexuous since
supporting structures are absent.
REMARKS. The surface reticulation, the absence of any tabulation and the apical
archaeopyle together distinguished this species from all previously described forms.
OCCURRENCE. C. reticulata is rare to common in samples from the Middle and
Upper Cenomanian of Fetcham Mill (not found below sample FM 750), and from
the Lower, Middle and Upper Cenomanian of Escalles (not found below sample
E 207). This species was absent from the samples from Compton Bay and from
North America.
Genus EPELIDOSPHAERIDIA nov.
DERIVATION OF NAME. Greek, epelidos, cover or lid; sphaera, ball — with reference
to the conical apical operculum which sometimes remains attached to the shell.
DIAGNOSIS. Shell subpolygonal; epitract conical with small apical protuberance,
hypotract polygonal with small antapical horn on one side. Shell wall two layered,
periphragm giving rise to a moderate number of spines, truncated or forked distally
Cingulum and sulcus outlined by spines. Cingulum slightly laevo-rotatory. Apical
archaeopyle.
TYPE SPECIES. Palaeoperidinium spinosum Cookson & Hughes 1964. Cambridge
Greensand (Cenomanian), England.
REMARKS. The presence of an apical archaeopyle, together with a well developed
cingulum and sulcus differentiate Epdidosphaeridia from all previously described
genera. Doidyx Sarjeant (19666) is most similar but differs in that the shell is asym-
metrical, the hypotract is conical and a sulcus is absent.
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i 143
Epelidosphaeridia spinosa (Cookson & Hughes) comb. nov.
(PI. 3, figs. 10-12)
1964 Palaeoperidinium spinosum Cookson & Hughes : 49, pi. 8, figs. 6-8.
1967 Palaeoperidinium spinosum Cookson & Hughes: Clarke & Verdier: 70, pi. 14, figs. 10-12.
DESCRIPTION. The shell possesses convex sides, conical epitract and a hypotract
which is more or less truncated posteriorly. A small apical prominence is commonly
present. The periphragm is smooth or lightly granular and forms a moderate num-
ber of small, stout spines. The spines appear to be hollow, closed proximally,
parallel sided, and are oblate distally or terminate with a small fork. The spines
widen slightly before joining the shell and they are sometimes joined proximally.
This is particularly well developed in the antapical region where the processes are
joined medially and form a slight projection on one side of the shell. The cingulum
is clearly delimited by two parallel lines of closely set spines and is 5 to 8 p in width.
It is only slightly helicoid and bears few spines on its surface. The sulcus in most
specimens is clearly defined, being slightly hollowed and almost devoid of spines.
The spines on the remainder of the shell surface are usually randomly arranged, but
occasionally a vague alignment is present suggesting a tabulation. An apical archa-
eopyle is constantly developed, the margin being only slightly angular.
DIMENSIONS. Range of observed specimens: shell length 32 (43) 56 fj,, width
27 (42-1) 57 p, maximum length of spines 2-5 (3-7) 5ju,. Number of specimens
measured, 21.
REMARKS. The Cenomanian specimens examined strongly resemble the type
material from the Cambridge Greensand and Chalk Marl (Lower Cenomanian) of
Cambridgeshire, England. In the lower horizons of the Cenomanian the sulcus,
although always present, is not so clearly defined as in higher horizons.
OCCURRENCE. E. spinosa is a rare to common species in the lower and middle
horizons of the Cenomanian of Fetcham Mill, Compton Bay and Escalles. It is
absent from samples FM 690, 670, 650; CB 13, 15, 17, 19, 21; and E 159, 153. It
has only beeen observed in one North American sample — sample Sas 1023
(Saskatchewan, Albian).
Cyst-Family HYSTRICHOSPHAERIDIACEAE Evit emend.
Sarjeant & Downie 1966
Genus HYSTRICHOSPHAERIDIUM Deflandre emend. Davey & Williams 1966
Hystrichosphaeridium tubiferum (Ehrenberg)
(PI. 5, figs. 5, 8)
1838 Xanthidium tubiferum Ehrenberg : pi. i, fig. 16.
19666 Hystrichosphaeridium tubiferum (Ehr.) Davey & Williams : 56, pi. 6, figs, i, 2; pi. 8, fig. 5;
pi. 10, fig. 2; text-fig. 13. (See also for earlier references).
DIMENSIONS. Range of observed specimens: diameter of central body 28 (38-1)
51 /z, maximum length of processes 15 (25-0) 37 //,. Number of specimens measured,
26.
i44 CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
REMARKS. H. tubiferum is an infrequent to common species at all horizons
throughout the Cenomanian of Fetcham Mill, Compton Bay and Escalles. It is also
recorded from the Albian (sample FM 886) and Turanian (sample FM 520) of Fetcham
Mill. This species was not recorded in the North American material. Thus the
earliest recording of H . tubiferum is from the Albian ; it ranges throughout the Upper
Cretaceous and has been recorded from the Eocene (Ypresian) by Davey & Williams
(19666). It is a long-ranging species of little stratigraphic value.
Hystrichosphaeridium deanei Davey & Williams
(PI. 4, %. i)
19666 Hystrichosphaeridium deanei Davey & Williams : 58, pi. 6, figs. 4, 8.
1967 Hystrichosphaeridium stellatum Maier; Clarke & Verdier: 55, pi. 12, figs, i, 2.
DIMENSIONS. Range of observed specimens: diameter of central body 37 (45-7)
54 fji, maximum length of processes 22 (35-8) 45 [i. Number of specimens measured,
10.
REMARKS. One specimen, occurring in sample E 207 (Escalles), possesses broad
processes and appears to occupy a position midway between H. deanei and H.
tubiferum.
OCCURRENCE. H. deanei is a rare species confined to the Middle and Upper
Cenomanian of Fetcham Mill, Compton Bay and Escalles (Table 25). The samples
in which it first occurs are FM 710 (Fetcham Mill), CB 13 (Compton Bay) and E 195
(Escalles).//. deanei has also been recorded from the Turonian sample FM520,
from Fetcham Mill. It is absent from the North American material.
Hystrichosphaeridium readei Davey & Williams
19666 Hystrichosphaeridium readei Davey & Williams : 64, pi. 6, fig. 3 (See also for earlier
references) .
DIMENSIONS. Range of observed specimens: diameter of central body 31 (42-1)
57 p, maximum length of processes 23 (29-7) 35 /t. Number of specimens measured,
ii.
OCCURRENCE. H. readei is very rare in samples FM 810, 790, 770 (Fetcham Mill)
and E 207 (Escalles); and it is rare to infrequent in samples FM 690, 670, E 183, 177,
X65, 159 and CB 9 (Compton Bay). Two specimens were recorded in the Albian
sample (FM 886) from Fetcham Mill. This species was not recorded in the North
American samples.
Hystrichosphaeridium radiculatum Davey & Williams
(PI. 4, fig- 8)
19666 Hystrichosphaeridium radiculatum Davey & Williams : 65, pi. 7, fig. 9; pi. 9, fig. 6.
DIMENSIONS. Range of observed specimens: diameter of central body 27 (38-8)
43 ju, maximum length of processes 12 (15-9) 20 /*. Number of specimens measured,
II.
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 145
OCCURRENCE. H. radiculatum is a very rare to infrequent species occurring at a
number of horizons throughout the Cenomanian of Fetcham Mill, Compton Bay and
Escalles. It has also been recorded in the Albian sample from Fetcham Mill
(sample FM 886), but is absent from the North American material.
Hystrichosphaeridium mantelli Davey & Williams
(PI- 4. ng. 9)
19666 Hystrichosphaeridium mantelli Davey & Williams : 66, pi. 6, fig. 6.
DIMENSIONS. Range of observed specimens: diameter of central body 32 (38-6)
48 fi, maximum length of processes 12 (21-4) 26^. Number of specimens measured,
15-
REMARKS. The reticulate nature of the central body and the fibrous processes
differentiate H. mantelli from most previously described species. H. radiculatum is
the most similar but differs from H. mantelli by the more branched and deeply
furcate processes and the tendency for the fibrils of the processes to continue across
the surface of the central body.
OCCURRENCE. H. mantelli is a very rare to infrequent species occurring in most
samples throughout the Cenomanian of Fetcham Mill, Compton Bay and Escalles.
It has also been recorded in the Turonian sample from Fetcham Mill (sample FM
520), but is absent from the North American material.
Hystrichosphaeridium bowerbanki Davey & Williams
(PI- 5, fig- 9)
19666 Hystrichosphaeridium bowerbanki Davey & Williams : 69, pi. 8, figs. 1,4.
DIMENSIONS. Range of observed specimens: diameter of central body 25 (31-9)
40jLt, maximum length of processes 20 (25-5) 28 p. Number of specimens measured,
10.
REMARKS. H. bowerbanki is rare to infrequent in six Middle Cenomanian samples —
FM 770, 750, 730 and 690 from Fetcham Mill and CB 9, and 17 from Compton
Bay. It has also been recorded from the Albian sample (FM 886) and the Turonian
sample (FM 520), both from Fetcham Mill.
Hystrichosphaeridium difficile Manum & Cookson
(PI. 4, figs. 2, 6, 7)
1964 Hystrichosphaeridium difficile Manum & Cookson : 12, pi. 3, figs. 1-3, 7.
DESCRIPTION. The shell is subspherical, sometimes with a small apical prom-
inence; shell wall thick (c. i /JL), smooth to lightly granular. The processes, approxi-
mately 30 in number, are complex, usually broadly tubiform or buccinate in shape,
of constant length on any specimen but variable in width (3 to 18/11). Distally the
146 CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
larger processes have a rectangular opening, with a serrate margin which gives rise
to four broad spines. Typically there are four bands of thickening extending along
the length of the processes and passing onto the shell surface. There, each joins
with a similar thickening from a neighbouring process, thus forming a coarse reticula-
tion on the shell surface. The large tubular processes are arranged in a circular
manner around the shell, reflecting the precingular, cingular and postcingular series
of plates. Fine processes are uncommon and may be sulcal in position. An apical
archaeopyle is typically developed and possesses an angular margin marked at
intervals with V-shaped notches. Although often remaining attached, isolated
operculae have been identified (PI. 4, figs. 6, 7) and bear four moderate-sized tubular
processes.
DIMENSIONS. Range of observed specimens: diameter of central body 49 (64-9)
79 p,, maximum length of processes 18 (24-5) 31 p. Number of specimens measured,
8.
REMARKS. The specimens studied are extremely similar to the type material of
Lower Cretaceous age described by Manum & Cookson (1964) from Arctic Canada.
The only difference appears to be that in the type material the shell surface has a
fine reticulation which is absent from the specimens studied.
Two similar species are H. costatum Davey & Williams (19666) from the Oxford
Clay of England and H. readei Davey & Williams (19666) from the Cenomanian of
England. However, both species are considerably smaller than H. difficile and possess
narrower and less complex processes.
OCCURRENCE. H. difficile has only been recorded from the Saskatchewan material.
It is infrequent in samples Sas 1084 (Albian) and Sas 890 (Cenomanian), and common
in sample Sas 805 (Cenomanian).
Genus OLIGOSPHAERIDIUM Davey & Williams 1966
Oligosphaeridium complex (White)
(PI. 5, figs. 6, 7)
1842 Xanthidium tubiferum complex White : 39, pi. 4, div. 3, fig. n.
19666 Oligosphaeridium complex (White) Davey & Williams : 71, pi. 7, figs, i, 2; pi. 10, fig. 3;
text-fig. 14 (See also for earlier references).
1967 Hystrichosphaeridium complex (White) Clarke & Verdier: 53, pi. n, figs. 10, n.
DIMENSIONS. Range of observed specimens: diameter of central body 34 (41-1)
55 /A, maximum length of processes 22 (34*6) 43 //,. Number of specimens measured,
12.
REMARKS. Davey & Williams described examples of 0. complex from the Speeton
Clay (Barremian), Lower Chalk (Cenomanian) and London Clay (Ypresian), all
from England.
OCCURRENCE. 0. complex is a rare to common species in all samples from Fetcham
Mill, Compton Bay and Escalles. It is also present in the Albian sample (FM 886)
and the Turonian sample (FM 520) both from Fetcham Mill. This species is present
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 147
in the Albian and the lowermost Cenomanian sample from Saskatchewan — Sas 1084
1023, 967, and 890 — but is absent from the Texas samples.
0. complex thus has a known stratigraphic range from the Neocomian (Gocht
1959; Cookson & Eisenack 1958) to the Eocene, Ypresian (Davey & Williams 19666).
Oligosphaeridium reticulatum Davey & Williams
19666 Oligosphaeridium reticulatum Davey & Williams : 74, pi. 7, fig. 10.
DIMENSIONS. Range of observed specimens: diameter of central body 29-47/4,
length of processes 14-26/4. Number of specimens measured, 5.
OCCURRENCE. 0. reticulatum is an infrequent species occurring in the two lower
samples from Fetcham Mill, samples FM 840 and 810.
Oligosphaeridium prolixispinosum Davey & Williams
(PI- 5, %. 4)
19666 Oligosphaeridium prolixispinosum Davey & Williams : 76, pi. 8, figs. 2, 3.
DIMENSIONS. Range of observed specimens: length of central body 33 (39-0)
43 /A, width 20 (28-7) 34 /u, maximum length of processes 18 (25-0) 30/4. Number of
specimens measured, 15.
OCCURRENCE. 0. prolixispinosum is a rare species confined mainly to the Lower
and Middle Cenomanian of Fetcham Mill and Escalles, although it does occur occas-
ionally in the Upper Cenomanian. This species has only been recorded twice in the
samples CB 17 and 21 from Compton Bay, both from the Upper Cenomanian. It
has not been recorded in the North American material.
Oligosphaeridium anthophorum (Cookson & Eisenack)
(PI. 5, figs, i, 2, 3)
1958 Hystrichosphaeridium anthophorum Cookson & Eisenack : 43, pi. n, figs. 12, 13; text-figs.
16-18.
1958 Hystrichosphaeridium anthophorum Cookson & Eisenack; Eisenack, 402, pi. 26, figs, i, 2.
1961 Hystrichosphaeridium anthophorum Cookson & Eisenack; Alberti : 34, pi. 9, fig. 16.
19666 Oligosphaeridium anthophorum (Cookson & Eisenack) Davey & Williams : 77.
DESCRIPTION. The shell is subspherical ; shell wall smooth to lightly granular.
An apical archaeopyle is typically developed and possesses an angular margin. The
processes are hollow, buccinate to infundibular, with the distal flared portion per-
forate. Distally the margins of the processes are usually entire, but may bear one
or two small spines. A complete specimen possesses 18 processes, reflecting a
tabulation characteristic of this genus. The processes are all of equal size except
for the first postcingular (i"') and the posterior intercalary (ip) which are often
reduced.
DIMENSIONS. Range of observed specimens: diameter of central body 38 (46-3)
57 /z, maximum length of processes 18 (34-1) 43 /*. Number of specimens measured, 8.
148 CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i
REMARKS. The presence of complex perforate processes having an entire distal
margin differentiate this species from all other similar forms.
OCCURRENCE. 0. anthophorum has been recorded from the Upper Jurassic —
Lower Cretaceous (Aptian-Albian) of Australia (Cookson & Eisenack 1958), from
the Aptian of Germany (Eisenack 1958) and from the Upper Barremian — Albian of
Germany (Alberti 1961). This species has been recorded from one sample, Sas 1023,
from the Albian of Saskatchewan where it is common.
Oligosphaeridium reniforme (Tasch)
(PI. 6, fig. i)
1964 Hystrichosphaeridium reniforme Tasch : 193, pi. 2, fig. 6.
19666 Oligosphaeridium reniforme (Tasch) Davey & Williams : 77.
DESCRIPTION. The shell is subspherical to ovoidal, shell wall lightly granular.
The processes are hollow, tubiform, widening distally into a broad, flat-topped funnel.
The distal margin of the funnel bears a small number of pointed and irregularly
shaped spines. Distally the processes sometimes possess large, subcircular perfora-
tions. An apical archaeopyle is typically developed.
DIMENSIONS. Range of observed specimens: diameter of central body 31 (42-4)
49 fj,, maximum length of processes 20 (24-9) 30 p. Number of specimens measured, 7.
REMARKS. The specimens appear to be very similar to the type material from
the Albian of Kansas although Tasch did not describe the presence of distal perfora-
tions. 0. reniforme differs from 0. anthophorum and 0. perforatum (Gocht 1959) by
the presence of distal spines. It differs from 0. pulcherrimum Deflandre & Cookson
(1955) in that the processes possess only a few spines, are not so complexly perforate
and distally are flat-topped.
OCCURRENCE. 0. reniforme is infrequent in all the Albian-Cenomanian samples
from Saskatchewan. It has not been recorded elsewhere.
Genus LITOSPHAERIDIUM Davey & Williams 1966
Litosphaeridium siphoniphorum (Cookson & Eisenack)
(PI. 6, figs. 3, 4; Fig. 15)
1958 Hystrichosphaeridium siphoniphorum Cookson & Eisenack : 44, pi. n, figs. 8-10.
19666 Litosphaeridium siphoniphorum (Cookson & Eisenack) Davey & Williams : 80, pi. 7,
figs. 7, 8; text-figs. 16, 17. (See also for earlier references).
1967 Hystrichosphaeridium siphoniphorum Cookson & Eisenack; Clarke & Verdier: 55, pi. n,
figs, i, 2.
DIMENSIONS. Range of observed specimens: diameter of central body 21 (34-0)
47 /u,, length of processes 4-25 //,. Number of specimens measured, 160.
DESCRIPTION. A statistical study was made of this species using 20-30 specimens
from each of six samples at 4O-foot intervals from the Fetcham Mill Borehole. The
purpose of this study was to see if the variation in the mean size of L. siphoniphorum
was directional, and if there was a significant difference in this measurement for
0)
-Q
E
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
149
40
30
B
50
60
50
60
70
40
50
60
70
40
50
60
40
50
60
40
50
60
70
Overall diameter
FIG. 15. Overall diameter — frequency histograms of Litosphaeridium siphoniphorum
(Cookson and Eisenack) at six horizons from Fetcham Mill, Surrey. A. Sample FM 650,
B. Sample FM 690, C. Sample FM 730, D. Sample FM 770, E. Sample FM 810, F. Sample
FM 840.
150
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
successive samples. The measurement taken was the overall diameter. The
position of the archaeopyle is always obvious, allowing easy specimen orientation.
Thus to make all measurements strictly comparable the overall diameter was always
taken in the plane of the archaeopyle.
Histograms (Fig. 15) were drawn for each assemblage. The mean overall diameter
for each assemblage varied for each horizon but, unfortunately, the variation was
not directional and, therefore, was of little stratigraphic value. The Student's
t-test was performed on successive pairs of assemblages to see whether or not they were
significantly different (Table A). A probability of 0-05 or less was taken as being
significant.
FM650
FM6go
FM6go
FM730
FM 770
FM 770
FMSio
FMSio
FM84o
(x= 50-0 ft)
x =
(x= 56-6 fj)
(x=48-3yM)
(x = 48-3/0
(x= 48-10)
(X = 48-I ft)
(X= 48-1 ft)
(x= 48-1/0
(x= 58-0/0
TABLE A
t = 3-08
t = 3-16
t = 0-084
t = o
t=5'4
(significant difference)
(significant difference)
(no significant difference)
(no significant difference)
(significant difference)
The results show that there is a significant difference at the 5 % level between
some of the assemblages with respect to this character. However, all the specimens
measured were apparently morphologically identical and differ only in size. Speci-
mens from one sample vary considerably in size but were probably formed by one
species of motile dinoflagellate. Thus the size of L. siphoniphorum appears to be
quite variable and should, at the moment, not be used as a diagnostic feature for the
subdivision of this species. The reason for the means in successive samples to be
significantly different is probably because of palaeoecological changes in the
environment.
REMARKS. All the specimens of L. siphoniphorum examined agree fairly closely
with the type material from Australia. The Surrey specimens appear to be smaller,
but the range of the Australian forms was not given so no true size comparison can
be made.
OCCURRENCE. L. siphoniphorum is rare to common at all horizons throughout the
Cenomanian of Fetcham Mill, Compton Bay and Escalles. It is present in the Albian
sample FM 866, but absent from the Turonian sample, FM 520. It is rare to
infrequent in the lower four samples from Saskatchewan — samples Sas 1084, 1023, 967
and 890. This species is also present in the Upper Woodbine Formation of Texas.
Thus L. siphoniphorum has a wide geographical distribution and a fairly restricted
range. It has been recorded from the Albian of Australia, Rumania, Canada and
Britain and from the Cenomanian of Australia, Canada and Britain.
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i 151
Genus POLYSPHAERIDIUM Davey & Williams 1966
Polysphaeridium pumilum Davey & Williams
?I955 Hystrichosphaeridium recurvatum White; Deflandre & Cookson : 269, pi. i, fig. 12.
19666 Polysphaeridium pumilum Davey & Williams : 93, pi. 7, figs. 3, 4.
DIMENSIONS. Range of observed specimens : overall diameter 30-40 //,, diameter
of central body 17-25/1,, length of processes 7-10/1,, width of processes 1-1-5 /A,
number of processes 38-44. Number of specimens measured, 3.
OCCURRENCE. Only three specimens of P. pumilum have been observed, one
from sample FM 750 and two from sample FM 770.
Polysphaeridium laminaspinosum Davey & Williams
(PL 4, figs. 10, ii)
19666 Polysphaeridium laminaspinosum Davey & Williams : 94, pi. 8, fig. 8.
DIMENSIONS. Range of observed specimens: diameter of central body 20 (26-8)
29 [L, maximum length of processes 9 (13-7) 17 p. Number of specimens measured, 8.
OCCURRENCE. P. laminaspinosum is rare to very rare, occurring spasmodically
throughout the Cenomanian of Fetcham Mill, Compton Bay and Escalles. It has
not been recorded elsewhere.
Genus TANYOSPHAERIDIUM Davey & Williams 1966
Tanyosphaeridium variecalamum Davey & Williams
(PI. 6, figs. 2, 5)
19666 Tanyosphaeridium variecalamum Davey & Williams : 98, pi. 6, fig. 7; text-fig. 20.
DIMENSIONS. Range of observed specimens : length of central body 27 (32-3) 43 /*,
width 14 (20-1) 24 n, maximum length of processes 11 (15-0) 24/4. Number of
specimens measured, 14.
OCCURRENCE. T. variecalamum is a rare to infrequent species at most horizons
throughout the Cenomanian of Fetcham Mill, Compton Bay and Escalles. It is
also present in the Albian sample, FM 886, and the Turonian sample, FM 520,
both from Fetcham Mill. One specimen was located in the Saskatchewan material,
from the Albian sample Sas 1084.
Genus CALLAIOSPHAERIDIUM Davey & Williams 1966
REMARKS. Hexasphaera Clarke & Verdier (1967; 42) is a junior synonym of
Callaiosphaeridium .
GEOL. 17, 3 IX
152 CENOMANIAN NON-CALCAREOUS MICROPLAN KTON, i
Callaiosphaeridium asymmetricum (Deflandre & Courteville)
(PI. 6, fig. 6)
1939 Hystrichosphaeridium asymmetricum Deflandre & Courteville : 100, pi. 4, figs, i, 2.
19666 Callaiosphaeridium asymmetricum (Deflandre & Courteville) Davey & Williams : 104,
pi. 8, figs. 9, 10; pi. 9, fig. 2.
1967 Hexasphaera asymmetrica (Deflandre & Courtville) Clarke & Verdier: 43, pi. 7, figs. 1-3
text-fig. 17.
DIMENSIONS. Range of observed specimens: diameter of central body 37 (45-8)
58 ju, maximum length of cingular processes 10 (25-0) 32/1,. Number of specimens
measured, 6.
REMARKS. The author disagrees with the description of Clarke & Verdier (1967)
in that the archaeopyle is epitractal, not apical, and that the antapical plate is
five-sided, not six-sided, bearing a process at each corner. The position of the three
sutural crests separating the large plates has, however, been verified.
OCCURRENCE. C. asymmetricum is rare to infrequent at most horizons throughout
the Cenomanian of Fetcham Mill, Compton Bay and Escalles. It has been recorded
in the Albian sample FM 886 and the Turonian sample FM 520, both from Fetcham
Mill, but is absent from the North American material.
Genus CLEISTOSPHAERIDIUM Davey, Downie, Sarjeant & Williams 1966
Cleistosphaeridium heteracanthum (Deflandre & Cookson)
(PI. 7, fig- 8)
1955 Hystrichosphaeridium heteracanthum Deflandre & Cookson : 276, pi. 2, figs. 5, 6; text-figs.
40, 41.
1966 Cleistosphaeridium heteracanthum (Deflandre & Cookson) Davey, Downie, Sarjeant &
Williams : 168, pi. 2, figs. 6, 7 (See also for earlier references).
DIMENSIONS. Range of observed specimens: shell diameter 42 (53-2) 63^,
maximum length of processes 9 (13-8) 17 ju. Number of specimens measured, 9.
OCCURRENCE. C. heteracanthum is a rare species restricted to the Upper Ceno-
manian— samples FM 690, 670 and 650 from Fetcham Mill, samples CB 19 and 21
from Compton Bay and sample E 153 from Escalles. It was not recorded from North
America.
Cleistosphaeridium multifurcatum (Deflandre)
(PI. 8, figs. 7, 10)
1937 Hystrichosphaeridium multifurcatum Deflandre : 76, pi. 16, figs. 1-3.
1939 Hystrichosphaeridium multifurcatum Deflandre; Deflandre & Courteville : 102, pi. 3,
fig. 2.
1952 Hystrichosphaeridium multifurcatum Deflandre; W. Wetzel : 400, text-fig. 16.
1955 Hystrichosphaeridium multifurcatum Deflandre; Valensi : 588, pi. i, fig. 21; pi. 5, fig. 5.
1960 Baltisphaeridium multifurcatum (Deflandre) Klement : 59.
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 153
1963 Hystrichosphaeridium multifurcatum Deflandre; Gorka : 66, pi. 9, figs. 4-6; text-fig. 8,
fig. i.
1966 Cleistosphaeridium multifurcatum (Deflandre) Davey, Downie, Sarjeant & Williams: 170.
DIMENSIONS. Range of observed specimens: diameter of central body 39 (55-0)
68 fj,, maximum length of processes 8 (ii'2) 13 //.. Number of specimens measured, 8.
OCCURRENCE. C. multifurcatum is rare at most horizons in the Cenomanian of
Fetcham Mill, Compton Bay and Escalles. In a few Upper Cenomanian samples
(FM 690, 670 and 650; CB 19, 21 and E 153) it is completely absent. In these
samples it seems to be replaced by C. heteracanthum. C. multifurcatum was not
observed in the North American samples.
Cleistosphaeridium armatum (Deflandre) comb. nov.
(PL 8, figs, i, 2, 12)
1937 Hystrichosphaeridium armatum Deflandre : 76, pi. 16, figs. 6, 7.
1947 Hystrichosphaeridium armatum Deflandre; Deflandre: fig. i, No. 10.
1952^ Hystrichosphaeridium armatum Deflandre; Deflandre: fig. 14.
1963 Baltisphaeridium armatum (Deflandre) Downie & Sarjeant : 91.
1967 Baltisphaeridium armatum (Deflandre) Clark & Verdier: 71, pi. 13, fig. 3.
EMENDED DIAGNOSIS. Shell subspherical ; shell wall of moderate thickness, densely
granular. Processes numerous, fairly broad, rigid, tapering gradually distally.
Proximally processes possess longitudinal basal striations; distally simple or giving
rise to variable number of small spines. Apical archaeopyle occasionally developed.
HOLOTYPE. Slide AJ. 54, Laboratoire de Micropaleontologie, Ecole Practique des
Hautes Etudes, Paris. (Figured by Deflandre 1937, pi. 16, fig. 6). Upper Cretaceous
flint from the Paris Basin.
DIMENSIONS. Holotype: shell length 20/11, width 18-20 /z, length of processes
10-15 p. Range of Cenomanian specimens: diameter of central body 19 (30-8) 42 /u,,
maximum length of processes 5 (9-6) 16 //,. Number of specimens measured, 33.
DESCRIPTION. The granules are elongate (c. 0-1-0-5 P in height) so giving the
shell surface the appearance of possessing a matting of short hairs. This ornamen-
tation, commented on by Deflandre in the original description of this species, was
verified by the present author when examining the type material in Paris.
The processes are all of a similar length on any one specimen and may vary in
width from i to 3 JM. They are hollow, always closed distally terminating either
simply (the extremity may be recurved) or more commonly the distal one quarter
of the processes bear a small number of stiff spines (pi. 8, fig. 2). The opening, when
observable, possesses an angular margin characteristic of an apical archaeopyle.
REMARKS. The specimens studied strongly resemble C. armatum as described
by Deflandre from Upper Cretaceous flints, one of which was of Cenomanian age.
The appearance of the shell surface and form of the processes make this an easily
recognizable species.
OCCURRENCE. C. armatum is common at all horizons throughout the Cenomanian
of Fetcham Mill, Compton Bay and Escalles. It is also present in the Upper Wood-
154 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
bine and Lower Eagle Ford formations of Texas, but has not been observed in the
Saskatchewan samples.
Cleistosphaeridium polypes (Cookson & Eisenack) comb. nov.
(PI. 6, figs. 7, 8)
19626 Hystrichosphaeridium recurvatum subsp. polypes Cookson & Eisenack : 491, pi. 4, figs.
11-13.
DESCRIPTION. The shell is spherical to subspherical and bears numerous slender
processes. The shell wall is thin and may be smooth or slightly granular. The
processes are narrow, usually less than i p in width for most of their length, broaden-
ing slightly proximally. They are closed distally and terminate in a number of
short, fine spines. The latter are not so numerous in the English and French speci-
mens as in those from Australia and North America where the distal spines are
numerous. A fairly large archaeopyle, probably apical, with an angular margin
is often present.
DIMENSIONS. Range of observed specimens : shell diameter 31 (38-4) 47 //,, maxi-
mum length of processes 9 (13-4) 18 /A. Number of specimens measured, 20.
REMARKS. C. polypes was originally described by Cookson & Eisenack from the
Aptian-Cenomanian of Australia and was considered to be a subspecies of
Hystrichosphaeridium recurvatum (White) . The resemblance between the two forms is
very slight and they are not considered to be closely related. C. polypes is placed
in this genus because of the presence of numerous closed processes and the probable
apical location of the archaeopyle.
OCCURRENCE. C. polypes is a very rare species restricted to the Middle and Upper
Cenomanian of Fetcham Mill (samples FM 750, 730), Compton Bay (samples CB 9,
17, 21) and Escalles (samples E 189, 177, 165). It is common in the Upper Wood-
bine formation of Texas, and is present in one sample from Saskatchewan (sample
Sas 835).
Cleistosphaeridium polypes var. clavulum nov.
(PI. 6, figs. 9, 10)
1964 Hystrichosphaeridium recurvatum subsp. polypes Cookson & Eisenack; Cookson & Hughes :
47, pi. 9, fig. 14.
DERIVATION OF NAME. Latin, clavulus, small nail — with reference to the pin-like
shape of the processes.
DIAGNOSIS. A variety of C. polypes possessing fine, capitate processes. Processes
terminating with fine spines are extremely rare.
TYPE. G.S.M. slide PF 3995(1). Lower Chalk, H.M. Geological Survey Borehole,
Fetcham Mill, Surrey at 840 feet depth. Upper Cretaceous (Cenomanian).
DIMENSIONS. Type: diameter of central body 29 by 32 /x, length of processes
12-13 /A. Range: diameter of central body 29 (31-5) 39 p, maximum length of
processes 13 (14-7) 15 p. Numbers of specimens measured, 10.
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON. i 155
REMARKS. C. polypes var. davulum differs from C. polypes in that the processes
are capitate, the terminal bulge being flattened and resembling the head of a pin.
Two specimens have been observed each possessing one process of the type charac-
teristic of C. polypes thus indicating a relationship between the two forms. C.
polypes probably evolved from this variety, spiny processes replacing the capitate
ones.
OCCURRENCE. This variety has only been recorded from the lowermost Cenoman-
ian samples at Fetcham Mill (sample FM 840) and Compton Bay (sample CB i).
It was previously recorded by Cookson & Hughes (1964) from the Upper Albian —
basal Cenomanian of England and so appears to be of stratigraphic importance in
England for indicating the base of this stage. It does not occur in the lowermost
sample from Escalles.
Cleistosphaeridium huguonioti (Valensi) comb. nov.
(PI. 7, fig. 10)
1955 Hystrichosphaeridium huguonioti Valensi : 38, fig. 2a.
ig6oa Hystrichosphaeridium ancoriferum Cookson & Eisenack : 8, pi. 2, fig. n.
1963 Hystrichosphaeridium ancoriferum Cookson & Eisenack; Baltes : 586, pi. 6, fig. 13.
1964 Hystrichosphaeridium ancoriferum Cookson & Eisenack; Cookson & Hughes : 47, pi. 9,
fig. 7-
1964 Chlamydophorella nyei Cookson & Eisenack; Cookson & Hughes : 54, pi. 6, fig. 12.
1966 Cleistosphaeridium ancoriferum (Cookson & Eisenack) Davey, Downie, Sarjeant &
Williams: 167, pi. 9, fig. i.
1967 Hystrichosphaeridium huguonioti Valensi; Clarke & Verdier: 54, pi. n, fig. 4, 5.
DESCRIPTION. The shell is subspherical; the shell wall is smooth and gives rise to
numerous bifurcating processes which are not aligned to any noticeable extent.
Most of the specimens possess an apical archaeopyle, the shape of which is usually
difficult to determine because of distortion. However, detached apical regions are
common and are 6-sided. The processes are hollow, the central cavity often being
constricted to some extent along its length, and closed distally and proximally.
The sides of the processes are practically parallel, diverging slightly proximally before
joining the shell. Distally they give rise to two slightly recurved spines. Cookson
& Eisenack comment on the " transparent tips " of the processes. The extremities
of the processes are in fact closed by a thin, transparent membrane. The processes
may be isolate or a few may be linked together distally by their spines. The shell
wall rarely forms a small rounded apical bulge.
DIMENSIONS. Range of observed specimens: diameter of central body 20 (31-8)
45 ju, length of processes up to 8 p. Number of specimens measured, 30.
REMARKS. Cookson & Hughes (1964) described C. huguonioti and Chlamydo-
phorella nyei from the Upper Albian and Lower Cenomanian of England, distinguish-
ing the latter, with difficulty, by the presence of an outer membrane and an apical
bulge. After a detailed examination of numerous Cenomanian specimens it was
concluded that Chlamydophorella nyei s.s., as described from Australia, does not occur in
the English and French Cenomanian and that C. nyei as described by Cookson &
156 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
Hughes belongs in Cleistosphaeridium huguonioti. C. huguonioti does occasionally
appear to possess a membrane linking some of the processes when their spines are
joined distally, and an apical prominence may also rarely be present. Some speci-
mens possess joined processes but apparently no apical prominence, whereas others
possess an apical prominence but no joined processes. The presence of distinctive
bifurcate processes and an apical prominence indicates that C. huguonioti is related
to Chlamydophorella, the outer membrane of the latter being reduced to a small,
transparent membrane at the distal end of each process.
OCCURRENCE. C. huguonioti is common throughout the Cenomanian of Fetcham
Mill, Compton Bay and Escalles (Fig. 22). It is absent from the uppermost sample
from Escalles (sample E 153) and from the Turonian sample FM 520. In the Upper
Cenomanian at these localities C. huguonioti tends to be replaced by its variety,
C. huguonioti var. pertusum nov., which appears to become more abundant as C.
huguonioti declines. It is present in the Albian sample FM 886 and is common in
the Upper Woodbine and Lower Eagle Ford formations of Texas, but was not
observed in the Saskatchewan material. This species was first recorded from the
Albian-Cenomanian of Australia and is also present in the Albian-Cenomanian
of Rumania (Balte§, 1963). Thus C. huguonioti appears to be wide-spread
geographically and restricted to the Albian and Cenomanian.
Cleistosphaeridium huguonioti var. pertusum nov.
(PL 7, figs. 6, 7, 9)
DERIVATION OF NAME. Latin, pertusum, perforated — with reference to the per-
forate appearance of the processes.
DIAGNOSIS. A variety of C. huguonioti possessing spherical to subspherical,
smooth walled shell bearing numerous processes. Processes broad-based, tapering
distally and terminating with two small recurved spines. Lumen of processes
restricted by transverse septa.
HOLOTYPE. G.S.M. slide PF 3040(2). Lower Chalk, H.M. Geological Survey
Borehole, Fetcham Mill, Surrey at 670 feet depth. Upper Cretaceous (Cenomanian).
DIMENSIONS. Holotype : diameter of central body 36 by 36 p,, length of processes
7-9 IJL, maximum width of processes distally 2 /*. Range : diameter of central body
24 (34-1) 46 //,, length of processes 6 (8-7) II p, maximum width of processes distally
i (2-7) 4-5 p. Number of specimens measured, 24.
DESCRIPTION. The processes taper distally from a fairly broad base (2-5-4^ in
width) to a narrow neck (c. 0-5 p in width) before bifurcating to give two short,
recurved spines. The processes are hollow but the lumen is traversed by a number
of small septa which thus subdivide it, giving the processes a " holey " or vacuolated
appearance. An archaeopyle is only rarely observable.
REMARKS. This variety differs from C. huguonioti in the form of its processes
which are vacuolated, tend to be longer and bear relatively small distal spines.
C. huguonioti found in the same samples possesses processes which are considerably
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i 157
wider distally (5-5-7-5 /i). C. huguonioti var. pertusum appears to have evolved from
C. huguonioti in the Middle to Upper Cenomanian.
OCCURRENCE. C. huguonioti var. pertusum occurs only in the Upper Cenomanian,
where it is common at all horizons. It first occurs in samples FM 710 (Fetcham Mill),
CB 15 (Compton Bay) and E 171 (Escalles). At these horizons it is occasionally
difficult to distinguish from C. huguonioti. Like C. huguonioti, this variety is absent
from sample E 153 (Escalles) and FM 520 (Fetcham Mill, Turonian). It has not
been observed in the samples from North America.
fCleistosphaeridium flexuosum Davey, Downie, Sarjeant & Williams
(PI. 7, figs. 4)
1966 ? Cleistosphaeridium flexuosum Davey, Downie, Sarjeant & Williams: 169, pi. 2, fig. 5.
OCCURRENCE. This is a very rare species occurring at most horizons throughout
the Cenomanian of Fetcham Mill. It has not been recorded elsewhere.
? Cleistosphaeridium parvum sp. nov.
(PL 7, figs, ir, 12)
DERIVATION OF NAME. Latin, parvus, little — with reference to the small size of
this cyst.
DIAGNOSIS. Shell ovoidal, small; shell wall smooth, bearing numerous long, fine
spines. Spines may be aligned along upper and lower boundaries of cingulum.
Cingulum strongly laevo-rotatory. Apical archaeopyle typically developed.
HOLOTYPE. B.M. (N.H.) ¥.51981 (3). Lower Chalk, Bureau de Recherches
Geologique et Minieres Borehole, Escalles, Pas de Calais at 165 metres depth. Upper
Cretaceous (Cenomanian) .
DIMENSIONS. Holotype: length of central body I2//,, width iiyu,, length of
processes 6-n p. Range: length of central body 11-14 /x, width 10-13 /z, length of
processes 6-12 /*. Number of specimens measured, 5.
DESCRIPTION. The spines are long, very fine, terminate distally in a point and
widen only slightly when joining the shell. The cingulum is not always observable
but some alignment of the spines parallel to the archaeopyle margin is usually pres-
ent.
REMARKS. ?C. parvum resembles only one previously described species, that is
Palaeostomocystis echinulata Deflandre (1937) from the Upper Cretaceous of France.
This species differs in that there are fewer spines (c. 12), but is similar in its ovoidal
shape, apical archaeopyle, cingulum and small size (6-7 //. long).
The presence of a cingulum has not previously been observed in members of
Cleistosphaeridium and might later be used as a character in generic subdivision.
However, difficulty of observation makes it of dubious value.
158 CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
OCCURRENCE. ?C. parvum is fairly common in three Upper Cenomanian samples
from Escalles — E 171, 165 and 159. It has also been observed in a single Lower
Cenomanian sample from Compton Bay (CB 5). In all other samples it appears to be
absent.
fCleistosphaeridium aciculare sp. nov.
(PL 6, figs, n, 12)
DERIVATION OF NAME. Latin, adcularis, like a needle — with reference to the
acuminate shape of the processes.
DIAGNOSIS. Shell spherical to subspherical ; shell wall of moderate thickness,
densely granular. Processes numerous, finely to broadly acuminate, slightly
flexuous, less than one-third of shell diameter in length.
HOLOTYPE. B.M. (N.H.) slide V. 51979 (3). Second White Speckled Shale,
International Yarbo Borehole No. 17, Saskatchewan at 835 feet depth. Upper
Cretaceous (Cenomanian).
DIMENSIONS. Holotype : diameter of central body 43 by 50 /A, length of processes
12-14 /*• Range: diameter of central body 32 (43-0) 54/^1, maximum length of
processes 8 (13-9) 21 //.. Number of specimens measured, n.
DESCRIPTION. The processes may be finely or broadly acuminate but on each
individual their width is constant. On individuals bearing fine processes these are
more densely packed than in individuals with broad processes. All intergradations
exist between the fine and the broad processed forms. The processes are always
pointed distally and occasionally bear small subsidiary spines near their extremities.
An archaeopyle has never been observed.
REMARKS. ?C. aciculare is only tentatively placed in this genus, for although it
resembles other members in overall appearance, an apical archaeopyle has not been
observed. The numerous acuminate processes and the densely granular shell sur-
face differentiate ?C. aciculare from most previously described forms. Most similar
seems to be Exochosphaeridium (Hystrichosphaeridiurri) cf. striolatum (Deflandre) as
illustrated by G6rka (1963, pi. 10, fig. 6) from the Cenomanian of Poland.
OCCURRENCE. 1C. aciculare is common in the following samples from Saskat-
chewan— Sas 1084, 1023 and 967 (all Albian) and Sas 835 (Cenomanian). It has
not been recorded elsewhere.
Genus SURCULOSPHAERIDIUM Davey, Downie, Sarjeant & Williams 1966
Surculosphaeridium longifurcatum (Firtion)
(PI. 8, fig. 9)
1952 Hystrichosphaeridium longifurcatum Firtion : 157, pi. 9, fig. i ; text-fig, i, H, K, L and M
1963 Baltisphaeridium longifurcatum (Firtion) Downie & Sarjeant: 91.
1966 Surculosphaeridium longifurcatum (Firtion) Davey, Downie, Sarjeant & Williams : 163,
pi. 8, figs. 7, ii ; text-figs. 43, 44.
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i 159
DIMENSIONS. Range of observed forms: lateral view — diameter of central body
30 (38-3) 47 /LI; apical view — diameter of central body 36 (42-2) 50/1., maximum
length of processes 14 (22-2) 29 /*. Mean diameter of archaeopyle, 20 //,. Number of
specimens measured, 24.
OCCURRENCE. This is a rare species at all horizons throughout the Cenomanian
of Fetcham Mill, Compton Bay and Escalles. The only exception is sample FM 730
from Fetcham Mill, where this species is very common, composing 22-5 % of the total
microplankton present. The reason for this unusual abundance is unknown. 5.
longifurcatum is also present in the Lower Eagle Ford formation of Texas, but was
not observed in the Saskatchewan samples. It has not been observed in the Albian
sample (FM 886) or the Turonian sample (FM 520) from Fetcham Mill, and thus
appears to be characteristically, Cenomanian.
Genus HYSTRICHOKOLPOMA Klumpp emend. Williams & Downie 1966
Hystrichokolpoma ferox (Deflandre)
(PL 9, figs. 5-7)
1937 Hystrichosphaeridium ferox Deflandre : 72, pi. 14, figs. 3, 4.
ig66a Hystrichokolpoma ferox (Deflandre) Williams & Downie : 181.
1967 Baltisphaeridium ferox (Deflandre) Clarke & Verdier: 73, pi. 15, fig. 4 (see also for earlier
references).
EMENDED DIAGNOSIS. Shell subspherical, densely granular or reticulate. Pro-
cesses thin walled, granular, often striated, of three kinds : (i) 6 large precingular and
4 large postcingular, possessing wide bases and tapering distally giving rise to 2 or
more tubules, typically open; (ii) a single long tubular antapical process and (iii)
cingular and sulcal processes, of moderate length, slender and tubular, only joining
proximally if at all. Archaeopyle apical.
HOLOTYPE. Slide AH 72, Laboratoire de Micropaleontologie, Ecole Practique
des Hautes Etudes, Paris. (Figured by Deflandre 1937^, pi. 14, fig. 3). Senonian
flint from the Paris Basin.
DIMENSIONS. Holotype: length of shell 46 p, width 36/4, overall length 78^,
length of processes 15-17 //.. Range of Cenomanian specimens : diameter of central
body 39 (46-1) 56 /z, maximum length of processes 27 (30-2) 36/11. Number of
specimens measured, 13.
DESCRIPTION. H. ferox appears to be a fairly variable species. In some speci-
mens the tabulation is not clearly indicated by the processes which tend to be smaller
and may even be closed distally. However, the processes usually are well developed,
their bases covering an area of the shell surface which is often slightly raised and
of the same shape as a thecal plate. The precingular and postcingular processes are
largest and give rise distally to as many as 10 tubules. Each cingular process
divides proximally into 2 or 3 long slender tubules aligned along the cingulum. The
sulcal processes may proximally divide into two tubules or may consist of a single
tubule which is sometimes reduced and closed distally. The antapical process is
long, tubular, open distally, and terminates with a smooth or serrate margin.
160 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
REMARKS. The Cenomanian specimens closely resemble the holotype of H . ferox,
which was examined in Paris by the author, by kind permission of Professor
Deflandre. The processes of the holotype are of the same form as, and similarly
positioned to, those of the Cenomanian forms. The antapical process of the holotype,
not shown in Deflandre's illustration (1937, pi. 14, fig. 3), is long and tubular.
OCCURRENCE. H . ferox is rare to very rare at most horizons throughout the Ceno-
manian of Fetcham Mill and Escalles, but has not been observed at Compton Bay
or in the North American samples. It is also present in the Turonian sample
(sample FM 520) from Fetcham Mill. Hence the stratigraphic range is from Aptian
(Eisenack 1958) to Upper Cretaceous, probably Senonian (Deflandre 1937).
Genus PROLIXOSPHAERIDIUM Davey, Downie, Sarjeant & Williams 1966
DIAGNOSIS. Shell elongate ovoidal to ellipsoidal, one pole (apical) typically lost
in archaeopyle formation. Opposite pole occupied by one or two antapical processes.
Remaining processes arranged in distinct rows, encircling shell and slightly offset
at a position corresponding to sulcus. Number of processes exceeds 30. Processes
closed proximally, typically but not constantly closed distally; their distal termina-
tions simple, faring in varied fashion, or briefly furcate. Shell surface sometimes
bears cover of coarse granules or very short, simple spinelets.
REMARKS. The diagnosis has been changed slightly: ' typically but not constantly
closed distally ' being inserted in place of ' closed or open distally ', with respect to the
processes. The processes of Prolixosphaeridium usually taper distally and are
closed. Specimens with open tubular processes belong to Tanyosphaeridium Davey
& Williams (19666).
Prolixosphaeridium conulum sp. nov.
(PI. 8, figs. 5, 6)
DERIVATION OF NAME. Latin, conulus, cone — with reference to the rather conical
shape of the processes.
DIAGNOSIS. Shell elongate ovoidal; shell wall densely granular and bearing
moderate number of processes. Processes acuminate to subconical, smooth walled,
typically rigid, pointed distally. Processes tend to be aligned in circular manner
around shell. Archaeopyle apical.
HOLOTYPE. B.M. (N.H.) ¥.51981 (5). Lower Chalk, Bureau de Recherches Geo-
logiques et Minieres Borehole, Escalles, Pas de Calais, at 165 metres depth. Upper
Cretaceous (Cenomanian).
DIMENSIONS. Holotype : shell length 47 //,, width 27 /u, length of processes 11-16 /*.
Range : shell length 38 (43-8) 50 \i, width 20 (25-9) 29 //,, maximum length of processes
ii (15-1) i8/i. Number of specimens measured, 8.
DESCRIPTION. The length of the shell is slightly less than twice the width. The
surface granules are relatively large (c. 0-2-0-4 /j, in width and height); they are
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 161
equidistant from one another. The processes (45-60 in number) have broad bases
(up to 6 fj,) and taper distally to terminate in a point. They are approximately
half the shell width in length, hollow and typically rigid, only occasionally being
bent near the distal end. In the central region of the shell the processes are aligned,
the rows encircling the shell. However, in one longitudinal portion in this region the
processes are generally smaller and haphazard in arrangement. This region probably
corresponds to the sulcus.
REMARKS. The distinctive shape, number and size of the processes distinguish
P. conulum. A rather similar but longer form was illustrated by Cookson & Eisenack
(1958, pi. 8, fig. IT ) as Hystrichosphaeridium parvispinum Deflandre. This specimen
comes from the Aptian of Australia and is said to grade into forms possessing more
numerous processes similar to P. conulum. Deflandre's species was placed in
Prolixosphaeridium by Davey, Downie, Sarjeant & Williams (1966).
Two other species are rather similar. The holotype of P. granulosum (Deflandre)
measures 18 by 33 /* and possesses 20-30 long processes, the latter being approxi-
mately equal to the shell width. The number of processes and the relative lengths
of the processes, therefore, differentiate this species from P. conulum. P. granulosum
as described by Valensi (1955) from the Upper Cretaceous, and Sarjeant (1962) from
the Upper Jurassic, are more similar to P. conulum but do not possess conical
processes.
P. mixtispinosum (Klement) differs from P. conulum by possessing two kinds of
processes — (i) approximately 50 processes of moderate length and (ii) numerous fine
hairs covering the shell surface.
OCCURRENCE. P. conulum has only been observed at certain horizons in the
Upper Cenomanian of Fetcham Mill, Compton Bay and Escalles — in samples FM 690,
CB 19, E 165, E 159 and E 153.
Genus CORONIFERA Cookson & Eisenack emend.
EMENDED DIAGNOSIS. Shell subspherical to ovoidal, bearing numerous simple or
bifurcating processes. Processes solid or hollow, closed distally, and joined proxi-
mally by low crests or membranes. Apical process distinctive, simple or branched.
Antapical process large, tubular, often terminating with denticulate margin. Archae-
opyle apical.
TYPE SPECIES. Coronifera oceanica Cookson & Eisenack 1958. Lower Cretaceous
(Albian) ; Australia.
REMARKS. The diagnosis has been emended to include the presence of an apical
archaeopyle and low crests joining the processes, and the positions of the two
distinctive processes.
Coronifera differs from Diphyes Cookson (1965) in that the processes are never
tubular and open distally, and by the presence of a reticulum joining the basal
portion of the processes. However, both genera possess a large tubular antapical
process and an apical archaeopyle, and are probably closely related.
162 CENOMANIAN NON -C ALC AREOU S MICROPL ANKTON, i
Coronifera oceanica Cookson & Eisenack
(PI. 8, figs. 8, ii)
1958 Coronifera oceanica Cookson & Eisenack : 45, pi. 12, figs. 5, 6.
1958 Coronifera oceanica Cookson & Eisenack; Eisenack : 407, pi. 25, fig. i.
1964 Coronifera oceanica Cookson & Eisenack; Cookson & Hughes : 56, pi. 9, figs. 8, 9.
1967 Coronifera oceanica Cookson & Eisenack; Clarke & Verdier: 77, pi. 17, fig. 7.
DESCRIPTION . The shell is subspherical to ovoidal, thin-walled, and bears numerous
processes of length between one-quarter and one-third of the shell diameter.
There is, occasionally, a slight apical prominence beneath the apical process. Fine
fibres radiate from the bases of the processes over the shell surface reminiscent of
those on the shell surface of Exochosphaeridium striolatum (Deflandre). The pro-
cesses are weak, fairly flexuous and commonly joined to each other by a network of
low crests or fine membranes. The latter may be proximal or may extend along the
entire length of the processes. Distally the processes are closed and may be simple,
bifurcate or trifurcate. A large tubular process is present at the antapex; it is open
distally and terminates with a denticulate margin. At the apex, when attached,
there is a process which is only slightly larger than the typical processes but is
usually branched and, therefore, distinctive. The large archaeoplye, developed in
the majority of specimens, has an angular margin and forms opposite the antapical
process.
A number of specimens of C. oceanica were observed in the Albian sample from
Fetcham Mill (FM 886). They resemble the specimens of Eisenack (1958) from the
Aptian of Germany in that the processes are fewer, more solid and are usually simple.
DIMENSIONS. Range of observed specimens: diameter of central body 31 (40)
54 /x, maximum length of processes 11 (15-3) 22 /z. Number of specimens measured, 15.
REMARKS. C. oceanica has previously been recorded from the Upper Aptian of
Germany (Eisenack, 1958), Albian of Australia (Cookson & Eisenack, 1958) and
basal Cenomanian of England (Cookson & Hughes, 1964). The surface reticulation
was not described in the Australian type material but appears to be present on the
photographed specimens. It was first commented on by Cookson & Hughes.
OCCURRENCE. In addition to the German and Australian records, C. oceanica is
infrequent to common at all horizons throughout the Cenomanian of Fetcham Mill,
Compton Bay and Escalles, and is also common in the Albian sample (FM 886) from
Fetcham Mill. It was not present in the Turonian sample (FM 520). One specimen
was observed in the North American material, in sample Sas 1084 (Albian) from
Saskatchewan. Thus the known stratigraphic range is from Upper Aptian to
Cenomanian.
Cyst-Family EXOCHOSPHAERIDIACEAE Sarjeant & Downie 1966
Genus EXOCHOSPHAERIDIUM Davey, Downie, Sarjeant & Williams 1966
REMARKS. Exochosphaeridium differs from Trichodinium Eisenack & Cookson
(1960) in that the latter possesses a well developed cingulum.
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i 163
Exochosphaeridium phragmites Davey, Downie, Sarjeant & Williams
(PI. 7> %. 5)
1966 Exochosphaeridium phragmites Davey, Downie, Sarjeant & Williams: 165, pi. 2, figs.
8-10.
DIMENSIONS. Range of observed specimens: maximum diameter of central
body 41 (51-9) 67 IJL, minimum diameter of central body 32 (46-7) 57 //,, maximum
length of processes 10 (18-5) 40 p,. Number of specimens measured, 18.
REMARKS. Superficially E. phragmites resembles E. striolatum (Deflandre) which,
however, has a definitely striated periphragm. Trichodinium paucispinum Eisenack
& Cookson (1960) is also similar but has fewer processes and a well developed
cingulum.
OCCURRENCE. E. phragmites is rare at most horizons throughout the Cenomanian
of Fetcham Mill, Compton Bay and Escalles, and is recorded from the Albian
sample, FM 886. It has not been recorded in the North American material.
Exochosphaeridium pseudohystrichodinium (Deflandre)
(PI. ii, figs. 4, 5)
1937 Hystrichosphaeridium pseudohystrichodinium Deflandre : 73, pi. 15, figs. 3, 4.
1966 ? Exochosphaeridium pseudohystrichodinium (Deflandre) ; Davey, Downie, Sarjeant &
Williams : 166.
1967 Baltisphaeridium pseudohystrichodinium (Deflandre); Clarke & Verdier: 75, pi. 15, fig. 7.
(see also for earlier references).
EMENDED DIAGNOSIS. Shell spherical to ovoidal; shell wall thick, with pitted
surface. Processes numerous, slightly fibrous, occasionally bifurcating medially,
slender, broadening slightly proximally ; distally truncated or terminated with small
bifurcation. Cingular processes rarely aligned. Apical process sometimes branched
and slightly larger than normal. Archaeopyle precingular, formed by loss of one
or two plate areas.
HOLOTYPE. Slide AH. 55, Laboratoire de Micropaleontologie, Ecole Practique
des Hautes Etudes, Paris. (Figured by Deflandre 1937, pi. 15, fig. 3). Upper
Cretaceous flint from the Paris Basin.
DIMENSIONS. Range of type material: shell length 49 to 54/11, shell width 38 to
45 /i, overall length 80-90 ju,. Range of observed specimens: diameter of central
body 35 (43-8) 54 p, maximum length of processes 13 (16-8) 21 /u,. Number of
specimens measured, 12.
DESCRIPTION. The shell wall is moderately thick (c. 1-5 ^), pitted (never striated),
and bears a large number of broad-based processes. All the processes appear to be
basically the same except for the apical process which is typically branched and is
usually larger than normal.
The author was permitted, through the courtesy of Professor Deflandre, to make a
detailed examination of the holotype and paratype. In the holotype the cingular
processes were arranged in a definite circular manner around the shell. Such an
164 CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
alignment was not observed with certainty in any of the Cenomanian specimens
studied. The archaeopyle, present in the paratype, is precingular and usually
formed by the loss of two plate areas.
REMARKS. The Cenomanian specimens differ from the holotype only in that
aligned circular processes were not observed. This may be due to unfavourable
preservation or orientation of the specimens. The diagnosis of E. pseudohystricho-
dinium has been emended to include a description of the apical process and the
archaeopyle.
The overall form of E. pseudohystrichodinium resembles species included in the
" hirsutum " group, but the typical fibrous shell periphragm is absent. The forma-
tion of the archaeopyle by the loss of two precingular plates has also been observed
in E. striolatum var. truncatum nov., indicating a relationship.
OCCURRENCE. This species is rather restricted, being common in samples FM 690,
E 165 and E 159 and rare in samples FM 750, E 207 and FM 520 (Turonian). It
appears, therefore, to be most common in the Upper Cenomanian. It was not
recorded in the samples from North America. The recorded stratigraphic range
is from Cenomanian to Eocene (Pastiels 1948).
Exochosphaeridium striolatum (Deflandre) comb. nov.
1937 Hystrichosphaeridium striolatum Deflandre : 72, pi. 15, figs, i, 2.
DIAGNOSIS. Shell subspherical to ovoidal. Processes numerous, variable,
fibrous, often bifurcate medially and sometimes terminating with small fork. Pro-
cesses distally may be pointed or blunted. Fibres pass down length of processes
onto shell surface and there join with similar fibres from adjacent processes. Apical
process and precingular archaeopyle may be present.
HOLOTYPE. Slide AH. 89, Laboratoire de Micropaleontologie, Ecole Practique
des Hautes Etudes, Paris. (Figured by Deflandre 1937, pi. 15, fig. i). Upper
Cretaceous flint from the Paris Basin.
REMARKS. Through the courtesy of Professor Deflandre, I was able to examine
the holotype and paratype of E. striolatum. On the holotype one process appeared
to be unusually thick and may have been apical in position. Neither holotype nor
paratype was observed to possess an archaeopyle but this may have been due to the
fact that the lower surfaces of the specimens were extremely dark and could not be
studied.
Exochosphaeridium striolatum (Deflandre) var. truncatum nov.
(PI. 7, figs. 1-3)
DERIVATION OF NAME. Latin, truncatus, shorten by cutting off — with reference
to the truncated extremities of the processes.
DIAGNOSIS. A variety of E. striolatum possessing subspherical to ovoidal shell;
shell wall fibrous or lightly pitted. Processes numerous, typically fibrous and blunted
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 165
distally, slender or subtriangular, rarely branched. Apical process large, often
foliate. Precingular archaeopyle, commonly present, formed by loss of two plate
areas, rarely one.
HOLOTYPE. B.M. (N.H.) ¥.51982 (i). Lower Chalk, Bureau de Recherches Geo-
logiques et Minieres Borehole, Escalles, Pas de Calais, at 159 metres depth. Upper
Cretaceous (Cenomanian).
DIMENSIONS. Holotype : diameter of central body 66 by 67 /z, length of processes
17-22^. Range: diameter of central body 34 (56-1) 81 ^, maximum length of
processes 6 (17-8) 27 p. Number of specimens measured, 24.
DESCRIPTION. The shell possesses a moderately thick wall (c. i /*) but it is quite
often distorted, especially when an archaeopyle is developed. The shell surface (peri-
phragm) is typically fibrous; the fibres pass down the length of the processes onto
the shell surface and, there, join up with similar fibres from adjacent processes.
Some specimens are less conspicuously fibrous, the fibres being apparent near the
bases of the processes and only extending a little way onto the shell surface. The
remainder of the shell surface in these forms is lightly pitted. The processes may be
fairly slender to subtriangular and are occasionally joined proximally. A small
number of processes are subdivided medially. The processes are typically trun-
cated distally but may be slightly bulbous. Process alignment was not observed.
The apical process is larger than the other processes and often foliate; the endo-
phragm occasionally forms a small apical bulge beneath it. The archaeopyle is
typically formed by the removal of two precingular plates, as is apparent by its shape.
Rarely only one plate is lost. Detached opercula consisting of two precingular
plates have been located (pi. 7, fig. 3).
REMARKS. E. striolatum var. truncatum nov. is a member of the " hirsutum "
group, a group which is characterized by having fibres radiating from the bases of
the processes over the shell surface. The truncate nature of the processes, although
obvious, has not previously been remarked on. Exochosphaeridium spinosum var.
deflandrei (Lejeuner-Carpentier 1941) is similar, the processes sometimes being
bulbous distally, but they are very slender and appear never to be truncated. Two
other forms, E. striolatum (Deflandre) (illustrated by Lejeune-Carpentier 1941 as
Hystrichosphaeridium hirsutum text-figs. 1-4) and E. (Hystrichosphaeridium) cf.
hirsutum (Cookson & Eisenack 1958), are also comparable, differing in that their
processes are not truncated. Baltisphaeridium bifidum Clarke & Verdier (1967)
is similar but possesses fewer and more slender processes which, however, are
bifid distally. The above forms all belong to the " hirsutum " group and are rather
similar, differing only in the detail form of the processes and perhaps in archaeopyle
formation, which is noted here for the first time.
Although the precingular archaeopyle is typically, but not always, formed by
the loss of two plate areas, this is not considered reason enough for the erection of a
new genus. This species may indicate a trend towards the genus Lingulodinium Wall
(1967) where four or five precingular plate areas are lost.
OCCURRENCE. E. striolatum var. truncatum is infrequent to fairly common at
all horizons throughout the Cenomanian of Fetcham Mill, Compton Bay and Escalles.
166 CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i
At Fetcham Mill it has also been recorded from the Albian (sample FM 886) and
Turonian (sample FM 520). A single specimen was obtained from the Upper Wood-
bine of Texas, but the species was absent from the Saskatchewan samples.
OTHER SPECIES
The following species and variety are here included in Exochosphaeridium on the
basis of similarity in structure :
Exochosphaeridium spinosum (White 1842) comb. nov. 1842 Xanthidium spinosum White:
Microsc. J., 11, 35-40, pi. 4 fig. 6.
Exochosphaeridium spinosum var deflandrei (Lejeune-Carpentier 1941) comb. nov. 1941 Hystrichos-
phaeridium spinosum var. defiandrei Lejeune-Carpentier: Annls. Soc. geol. Belg., 63 (bull, 3),
684, figs. 6, 7.
Cyst-Family AREOLIGERAGEAE Evitt emend. Sarjeant & Downie 1966
Genus CYCLONEPHELIUM Deflandre & Cookson emend. Williams & Downie 1966
REMARKS. Four species are described from the Cenomanian, C. distinctum Cook-
son & Eisenack being the only one fairly common throughout this stage. C. mem-
braniphorum Cookson & Eisenack, C. vannophorum sp. nov. and C. paucispinum sp.
nov. are restricted in occurrence. One species C. eisenacki sp. nov., is described from
the Albian of Saskatchewan. In the Cenomanian, the genus is most abundant at
Escalles, rarest at Saskatchewan and absent from Texas, possibly indicating a
preference for an open water environment.
Cyclonephelium distinctum Deflandre & Cookson
(PI. n, figs. 6-8, 10 ; Figs. 16 C, D, F)
1955 Cyclonephelium distinctum Deflandre & Cookson : 285, pi. 2, fig. 14.
1963 Circulodinium deflandrei Alberti, Baltes: 587, pi. 6, figs. 9-11.
1967 Cyclonephelium distinctum Deflandre & Cookson; Clarke & Verdier: 22, pi. i, figs. 6, 7.
(see also for earlier references) .
DESCRIPTION. This species is extremely variable, as pointed out by Cookson &
Eisenack (1962), and a member of most Cenomanian assemblages studied. The
shell, which is always somewhat flattened, may be subcircular to ovoidal in outline
and has a smooth or lightly granular shell wall. The regular outline is sometimes
broken by an apical protuberance and more rarely by two reduced, antapical horns.
The latter are of unequal size and when present the antapical region is slightly con-
cave (Fig. i6F). The bald areas, typical of this genus, may occupy almost all of the
ventral and dorsal surfaces of the shell leaving only the peripheral regions to bear
processes, or may be practically invisible beneath the encroaching processes.
Usually, however, these bald areas are circular to ovoidal in shape and occupy
approximately thirty per cent of each side of the shell. The processes are usually
abundant, extremely variable in form and typically under one-third of the shell
width in length. They are solid and usually widen distally and proximally, may be
fine or taeniate, and are often distally bifurcate. The bases of the processes are
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i 167
occasionally thickened and rarely a line of thickening on the shell surface joins one
process with a neighbouring one. Equally rarely, and only when the processes are
broad, they may be joined distally.
A large apical archaeopyle is always present. This is angular, possesses a pro-
nounced zigzag margin and on the ventral surface usually a relatively deep sulcal
notch. Although the archaeopyle is always discernible, the operculum is sometimes
still attached. It then behaved as a lid, returning to its original position after the
escape of the encysted organism.
DIMENSIONS. Range of observed specimens: height of shell (operculum absent)
41 (54-5) 82 ju, width 48 (61-8) 81 \L, maximum length of the processes 4 (10-7) 21 /z.
Number of specimens measured, 50.
REMARKS. C. distinctum appears to be extremely variable, and specimens showing
extremes in variation sometimes appear to be transitional to other species.
OCCURRENCE. Found throughout the Cenomanian of Fetcham Mill, Compton
Bay and Escalles, and rare to common at all horizons. Two samples from Sas-
katchewan, samples Sas 890 and Sas 805, also contain C. distinctum. The species
had a wide geographical distribution throughout most of the Cretaceous and is of
little value in detailed stratigraphy.
Cyclonephelium membraniphorum Cookson & Eisenack
(PL ii, fig. 9)
1958 Cyclonephelium compactum Deflandre & Cookson : 48, pi. 12, fig. 8 only.
19626 Cyclonephelium membraniphorum Cookson & Eisenack : 495, pi. 6, figs. 8-14.
1964 Cyclonephelium membraniphorum Cookson & Eisenack; Cookson & Hughes : 44, pi. 10,
figs. 5, 6.
1967 Cyclonephelium membraniphomm Cookson & Eisenack; Clarke & Verdier: 23, pi. 2,
figs, i, 2.
DESCRIPTION. The shell is always dorso-ventrally flattened, both the apex and
antapex being circular to subcircular in outline. An apical archaeopyle is always
developed, with a zigzag margin and sulcal notch. The bald areas, typical of this
genus, may be relatively large or quite small and are surrounded by high membranes.
The latter are braced by stout supports which arise from lines of thickening on the
shell surface. These lines of thickened periphragm may be curved and may form
semi-circular rings. When the latter are present, the membranes are in the form
of wide tubular projections. The periphragm of the shell wall and the membranes
is granular or pitted. The membranes may sometimes be fenestrate.
In some specimens attributed to C. membraniphorum the bald areas are practically
non-existent. The membranes in these forms are particularly well developed and
commonly are in the form of wide tubes or funnels. Distally they have a sub-
circular to polygonal outline and may possess a thickened outer rim bearing irregular
pieces of periphragm (PI. n, fig. 9).
DIMENSIONS. Range of observed specimens: length of shell (operculum missing)
37 (50-6) 79 ft, width 41 (54-1) 82 /u, maximum height of membranes 6 (13-3) 26^.
Number of specimens measured, 16.
GEOL. 17, 3 12
168 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
REMARKS. Many of the specimens observed appear identical with the Albian to
Cenomanian forms from Australia, except that the former are considerably smaller,
the largest of them only falling within the size range of the type material. It is
not thought practical to separate the European forms solely on the size distinction
and they have, therefore, been placed in the Australian species. An interesting
variation is the apparent absence of the bald areas, although the overall shape and
the apical archaeopyle typical of Cyclonephelium are present.
OCCURRENCE. In addition to the Australian records, C. membraniphorum is rare
and spasmodic throughout the Cenomanian of Fetcham Mill, Compton Bay and
Escalles, though absent elsewhere. This species has also been recorded from the
Turonian and Lower Senonian by Clarke & Verdier (1967).
Cyclonephelium vannophorum sp. nov.
(PL 9, fig. 3: PI. ii, figs, n, 12; Fig. i6E)
DERIVATION OF NAME. Latin, vannophorum, fan-bearer — with reference to the
fan-shaped processes.
DIAGNOSIS. Shell subcircular in outline, with slight apical prominence and one
or two reduced antapical horns. Shell wall coarsely granular with areas devoid of
processes of moderate size. Processes numerous, short, solid, of irregular shape and
often confluent distally. Apical archaeopyle with zigzag margin.
HOLOTYPE. B.M. (N.H.) slide ¥.51986 (i). Lower Chalk, Compton Bay, Isle of
Wight at 15 feet 6 inches above the base of the Cenomanian. Upper Creatceous
(Cenomanian) .
DIMENSIONS. Holotype: length of shell (including operculum) 65 /*, width 62 /u,
length of processes up to 8//,. Range: length of shell (without operculum) 47 (57-8)
70 /*, width 56 (65-5) 78 p, maximum length of processes 3 (5-3) 8/u,. Number of
specimens measured, 11.
DESCRIPTION. When two antapical horns are present the portion of the shell
between them is concave and from this region a broad furrow passes towards the
apex. The furrow or sulcus decreases in width and depth in this direction and
disappears just posterior to the archaeopyle margin. The processes vary greatly
in size from mere enlarged granules (0-5 ju,) to 8 p, in length. In the larger processes
the stem is quite narrow, the distal third widening rapidly (PI. 9, fig. 3) and is some-
times bifurcate. The processes are often joined to form a short line on the shell
surface. Rarely the cingulum is just discernible by a concentration of small
processes along its borders.
REMARKS. C. vannophorum sp. nov. is most closely comparable to ?C. attadalicum
Cookson & Eisenack (19626) from the Aptian /Albian of Australia. The processes
are similar in form but the shell of ?C. attadalicum is more polygonal and the cingulum
is always well defined.
OCCURRENCE . C . vannophorum has only been recorded from the Lower Cenomanian
sample CB 3 (Compton Bay) and it there comprises about 2% of the micro-
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
169
E F
FIG. 16. Cyclonephelium eisenacki sp. nov., A. Complete specimen (x 700). B. Specimen
illustrating apical archaeopyle ( x 700). Cyclonephelium distinctum Deflandre & Cookson,
C. D. and F. Illustrating variation in the shape of the shell of this species (processes re-
moved) (X 700). Cyclonephelium vannophorumsp. nov., E. Specimen with well developed
apical archaeopyle (processes removed) (X 700).
iyo CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
plankton content. The unusual distribution and the similarity to a Lower Cretaceous
species indicate that C. vannophorum is possibly a derived form.
Cyclonephelium paucispinum sp. nov.
(PI. 9, figs, i, 2)
DERIVATION OF NAME. Latin, paucus, few; spina, thorn — with reference to the
scarcity of the processes.
DIAGNOSIS. Shell subpolygonal in outline with well defined antapical horn.
Shell wall lightly to coarsely granular, bearing small number of irregularly distri-
buted processes confined to peripheral region of shell. Processes of moderate size,
solid and widening distally. Apical archaeopyle with a zigzag margin and sulcal
notch.
HOLOTYPE. B.M. (N.H.) ¥.51981 (2). Lower Chalk, Bureau de Recherches Geo-
logiques et Minieres Borehole, Escalles, Pas de Calais at 165 metres depth. Upper
Cretaceous (Cenomanian).
DIMENSIONS. Holotype : length of shell (operculum missing) 54 //,, width 81 p,
length of processes 1-5-12 /x. Range: length of shell (operculum missing) 40 (72-4)
92 /z, width 54 (87-0) 112 /A, maximum length of processes 5 (10-2) 19 p. Number of
specimens measured, 12.
DESCRIPTION. The shell is subpolygonal in outline, the cingular region being the
widest portion of the shell. Posteriorly there is a pointed antapical horn. The
few processes present are of moderate size and if close together tend to anastomose
both distally and proximally.
REMARKS. The large size, the paucity of processes and the subpolygonal shape
of the shell distinguish C. paucispinum sp. nov. from all previously described species.
The processes most closely resemble those of C. distinctum but are fewer in number.
OCCURRENCE. C. paucispinum is rare in samples E 165, CB 3, and common in
sample CB n. The distribution is, therefore, rather erratic in the Lower, Middle
and Upper Cenomanian.
Cyclonephelium eisenacki sp. nov.
(PI. 8, figs. 3, 4; PL 9, fig. 4; Figs. I7A, B)
1960 Aptea cf. polymorpha Eisenack & Cookson : 9, PI. 3, figs. 2-4.
DERIVATION OF NAME. The species is named after Professor A. Eisenack.
DIAGNOSIS. Shell subtriangular, flattened, with convex sides. Apical horn well
developed; antapical horns, if present, very reduced. Greater part of shell surface
ornamented by complex network of low crests and short, capitate processes. In
centre of both dorsal and ventral sides there is a circular area devoid of ornamenta-
tion. Apical archaeopyle always present.
CENOMANIAN NON-CALCAREOUS MICROPLANKTON, i 171
HOLOTYPE. B.M. (N.H.) V. 51980(2). Upper Lower Colorado (Fish Scale
Zone?), International Yarbo Borehole No. 17, Saskatchewan at 1023 feet depth.
Lower Cretaceous (Albian).
DIMENSIONS. Holotype: shell length 70 /z, width 66 p, height of crests 1-5 p.
Range: shell length 70-92/11 (4 complete specimens measured); shell length (oper-
culum missing) 55 (63-1) 71 /z, width 66 (72-1) 84 /z, height of crests 2 (4-2) 7 /z.
Number of specimens measured, 12.
DESCRIPTION. The shell surface is ornamented by a very characteristic network
of low crests or lamellae which are often broken, so forming isolated processes. A
poorly-marked cingulum, outlined by the crests, is occasionally discernible. The
apical operculum is often in position.
REMARKS. The illustrations of Aptea cf. polymorpha indicate that these speci-
mens belong in the genus Cydonephelium and may well be conspecific with the
Canadian specimens here described. This species differs from Aptea polymorpha
Eisenack (1958) by the absence of the numerous fine processes and the outer mem-
branous structure which they support. Pseudoceratium turneri Cookson & Eiesnack
(1958), from the Aptian /Albian of Australia, possesses a similar but stronger orna-
mentation. Also the apical and antapical horns are usually much better developed,
although the specimen of P. turneri illustrated (loc. cit. pi. 5, fig. 5), approaches a
number of the C. eisenacki examples.
OCCURRENCE. C. eisenacki sp. nov. is fairly common in two samples from
Saskatchewan, samples Sas 1084 and 1023. It has not been recorded elsewhere in
the material examined.
Cyst-Family ADNATOSPHAERIDIAGEAE Sarjeant & Downie 1966
Genus ADNATOSPHAERIDIUM Williams & Downie 1966
Adnatosphaeridium chonetum (Cookson & Eisenack) comb. nov.
(PL 10, figS. II, 12.)
19626 ICannosphaeropsis choneta Cookson & Eisenack : 493, pi. 4, figs. 8-10.
DESCRIPTION. The shell is spherical to subspherical and bears a number of more
or less tubular processes which possess bands of thickening for support. The pro-
cesses are usually joined to their neighbour either along their entire length, or only
distally by membranes. However, isolated tubular processes are occasionally
visible. The impression obtained from most specimens is of a complex membranous
network perpendicular to the shell surface. A well developed apical archae-
opyle is typically present.
DIMENSIONS. Range of observed specimens: shell diameter 27 (34-1) 46^, length
of processes 6 (10-4) 24 /z. Number of specimens measured, 15.
REMARKS. The Cenomanian specimens examined strongly resemble, but are
slightly smaller than, the type material from the Cenomanian of Australia. This
172 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
species, tentatively placed in Cannosphaeropsis by Cookson & Eisenack, is here
transferred to Adnatosphaeridium because of the presence of interconnecting
processes and an apical archaeopyle.
OCCURRENCE. Apart from the Australian occurrence, it is infrequent to common
in the Middle and Upper Cenomanian of Fetcham Mill and Escalles, not occurring
below samples FM 790 and E 195. Only one specimen was recorded from Compton
Bay, in sample CB 19.
Cyst-Family HYSTRICHOSPAERAGEAE O. Wetzel emend. Sarjeant
& Downie 1966
Genus HYSTRICHOSPHAERA O. Wetzel emend. Davey & Williams 1966
Hystrichosphaera ramosa var. ramosa (Ehrenberg)
(PI. 10, figS. I, 2, 5)
1838 Xanthidium ramosum Ehr. : pi. i, figs, i, 2, 5.
19660 Hystrichosphaera ramosa (Ehr.) var. ramosa Davey & Williams : 33, pi. i, figs, i, 6; pi. 3,
fig. i; text-fig. 8. (See also for earlier references).
1967 Hystrichosphaera furcata (Ehr.) Clarke & Verdier: 48, pi. 8, figs. 12, 13.
DIMENSIONS. Range of observed specimens: diameter of central body 29 (40-7)
56 fi, maximum length of processes 13 (19-8) 27 jii. Number of specimens measured,
32.
OCCURRENCE. This variety has a known stratigraphic range from the Oxfordian
(Jurassic) to the Ypresian (Eocene). It is a common variety throughout the Ceno-
manian of Fetcham Mill, Compton Bay and Escalles, and is rare to infrequent in the
material from Saskatchewan and Texas.
Hystrichosphaera ramosa (Ehrenberg) var. gracilis Davey & Williams
1955 Hystrichosphaera ramosa (Ehr.) Deflandre & Cookson : 263, pi. 5, fig. 8.
1963 Hystrichosphaera ramosa (Ehr.) Gorka : 48, pi. 6, figs. 6, 7.
1966^ Hystrichosphaera ramosa (Ehr.) var. gracilis Davey & Williams : 34, pi. i, fig. 5; pi. 5,
fig. 6.
DIMENSIONS. Range of observed specimens: diameter of central body 28-35 //,,
maximum length of processes 16-22 /u,. Number of specimens measured, 5.
OCCURRENCE. The known stratigraphic range of this variety is from the Ceno-
manian (England) to the Miocene (Australia). It is rare to infrequent at most hori-
zons throughout the Cenomanian of Fetcham Mill, Compton Bay and Escalles, but
has not been recorded in the North American material.
CENOMANIAN NON-CALCAREOUS MICROPLAN KTON, i 173
Hystrichosphaera ramosa (Ehrenberg) var. multibrevis Davey & Williams
(PL 10, figs. 3, 4)
I955 Hystrichosphaera furcata (Ehr.) Valensi : 586, pi. 4, fig. 4; pi. 5, fig. 12.
1958 Hystrichosphaera furcata (Ehr.) Eisenack : 406, pi. 25, figs. 4-8.
19660 Hystrichosphaera ramosa (Ehr.) var. multibrevis Davey & Williams : 35, pi. i, fig. 4;
pi. 4, fig. 6; text-fig. 9.
DIMENSIONS. Range of observed specimens : diameter of central body 31 (39-7)
51 n, maximum length of processes 10 (14-3) 21 p. Number of specimens measured, 13.
OCCURRENCE. H. ramosa var. multibrevis has a stratigraphic range from the
Lower Cretaceous (Hauterivian) to the Eocene (Ypresian). It is rare to infrequent
in all samples examined from Fetcham Mill, Compton Bay and Escalles. It has
also been recorded from the Lower Cretaceous (Albian) of Saskatchewan, in samples
Sas 967, 1023 and 1084.
Hystrichosphaera ramosa (Ehrenberg) var. reticulata Davey & Williams
19660 Hystrichosphaera ramosa (Ehr.) var. reticulata. Davey & Williams : 38, pi. i, fig. 2.
DIMENSIONS. Range of observed specimens: diameter of central body 33 (45-2)
59 p, maximum length of processes 13 (15-3) 17 //,. Number of specimens measured,
13-
OCCURRENCE. This variety is rare to infrequent at most horizons throughout the
Cenomanian of Fetcham Mill, Compton Bay and Escalles. It has also been
recorded in two Albian samples from Saskatchewan, samples Sas 967 and 1084.
Hystrichosphaera cingulata var. cingulata (O. Wetzel)
J933 Cymatiosphaera cingulata O. Wetzel : 28, pi. 4, fig. 10.
19660 Hystrichosphaera cingulata (O. Wetzel) Davey & Williams : 38, pi. i, fig. 9. (See also
for earlier references).
1967 Hystrichosphaera cingulata var. cingulata (O. Wetzel) Clarke & Verdier: 45, pi. 8, figs. 9, 10.
DIMENSIONS. Range of observed specimens: diameter of central body 26 (36-8)
48 ju, maximum height of crests 5 (7-0) 13 /z. Number of specimens measured, 16.
OCCURRENCE. H. cingulata is infrequent at all horizons throughout the Ceno-
manian of Fetcham Mill, Compton Bay and Escalles. It has not been observed in
the North American material. The recorded stratigraphic range is from Albian to
Middle Miocene.
Hystrichosphaera cingulata (O. Wetzel) var. reticulata Davey & Williams
19660 Hystrichosphaera cingulata var. reticulata Davey & Williams : 39, pi. i, fig. 10; pi. 2,
fig- 4-
1967 Hystrichosphaera cingulata var. perforata Clarke & Verdier : 46, pi. 9, figs. 2-4, text-fig. 19.
DIMENSIONS. Range of observed specimens: diameter of central body 33 (42-8)
59 ft, maximum height of crests 7 (10-2) 17 /u.. Number of specimens measured, 14.
174 CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i
OCCURRENCE. H. cingulata var. reticulata is rare to infrequent throughout the
Cenomanian of Fetcham Mill, Compton Bay and Escalles. It appears to be slightly
more common in the Middle and Upper Cenomanian at these localities. It has not
been observed in the North American samples. Clarke & Verdier (1967) also record
this variety from the Turonian and Senonian.
Hystrichosphaera crassimurata Davey & Williams
19663 Hystrichosphaera crassimurata Davey & Williams : 39, pi. i, fig. u.
1967 Hystrichosphaera cingulata var. polygonalis Clarke & Verdier: 47, pi. 8 figs. 7, 8, text-fig.
20.
DIMENSIONS. Range of observed specimens: diameter of central body 36-46 /z,
height of crests, up to 14 jit. Number of specimens measured, 4.
OCCURRENCE. H. crassimurata is very rare to rare in the Middle and Upper
Cenomanian of Fetcham Mill, Compton Bay and Escalles, the lowest samples in
which it is found being FM 790, E 195 and CB 9. It has also been recorded from the
Upper Woodbine Formation of Texas but is absent from the Saskatchewan material.
Hystrichosphaera crassipellis Deflandre & Cookson
1954 Hystrichosphaera crassipellis Deflandre & Cookson: text-fig. 5.
ig66a Hystrichosphaera crassipellis Deflandre & Cookson; Davey & Williams : 40, pi. i, figs. 7,
8. (See also for earlier references).
1967 Hystrichosphaera crassipellis Deflandre & Cookson; Clarke & Verdier: 48, pi. 8, fig. n;
pi. 9, fig. i.
DIMENSIONS. Range of observed specimens: diameter of central body 34 (47-9)
68 p,, maximum length of processes 10 (17-8) 29 p. Number of specimens measured, 19.
OCCURRENCE. H. crassipellis is rare at most horizons throughout the Cenomanian
of Fetcham Mill, Compton Bay and Escalles; it has not been recorded in the North
American material. The recorded stratigraphic range is from Cenomanian to Middle
Miocene (Gerlach 1961).
Genus ACHOMOSPHAERA Evitt 1963
Achomosphaera ramulifera (Deflandre)
(PL 10, fig. 7)
1935 Hystrichosphaera cf. ramosa (Ehr.) Deflandre : pi. 5, fig. n.
1966^ Achomosphaera ramulifera (Deflandre) Davey & Williams : 49, pi. 2, fig. 3. (See also for
earlier references) .
1967 Achomosphaera ramulifera (Deflandre); Clarke & Verdier: 40, pi. 8, fig. i.
DIMENSIONS. Range of observed specimens: diameter of central body 36 (44-7)
61 /A, maximum length of processes 16 (25-8) 36 p. Number of specimens measured, 14.
CENOMANIAN NON-CALCAREOUS MICROPL AN KTON, i 175
OCCURRENCE. This species has a known stratigraphic range from the Cenomanian
to the Eocene (Pastiels 1948). It is a very rare to infrequent species at most horizons
in the Cenomanian from Fetcham Mill, Compton Bay and Escalles. It is absent
from the North American samples.
Achomosphaera sagena Davey & Williams
ig66a Achomosphaera sagena Davey & Williams : 31, pi. 2, figs, i, 2.
1967 Achomosphaera reticulata Clarke & Verdier: 41, pi. 8, figs. 2, 3, text-fig. 16.
DIMENSIONS. Range of type material: diameter of central body 35 (48-4) 59^,
maximum length of processes 17 (20-8) 28 ^. Number of specimens measured, 12.
OCCURRENCE. A. sagena is a very rare to rare species occurring spasmodically in
the Cenomanian samples from Fetcham Mill, Compton Bay and Escalles. Clarke &
Verdier (1967) have also recorded it (as A . reticulata) from the Turonian and Senonian.
Genus Hystrichodinium Deflandre emend. Clarke & Verdier 1967
Hystrichodinium voigti (Alberti)
(PI. 10, figs. 6, 10)
1961 Heliodinium voigti Alberti : 33, pi. 8, figs. 1-5.
1966^ Heliodinium voigti Alberti; Sarjeant : 142, pi. 16, fig. 2; text-fig. 36.
1967 Hystrichodinium voigti (Alberti) Clarke & Verdier: 38.
DESCRIPTION. The shell is ovoidal to subpolygonal. The periphragm is smooth
or slightly granular, and forms sutural crests and ribbon-like processes. The former
are variably developed, may occasionally be absent, but when present give rise to
long, flexuous processes along their length. When the crests are absent the pro-
cesses arise directly from the shell surface. The processes are not confined to the
plate boundaries and may be distributed at random over the shell surface. The
processes are thin-walled, typically simple and occasionally terminate with bifurcate
or trifurcate extremities. The cingulum (4-5^ wide) is always delimited by low
crests and is strongly helicoid. A precingular archaeopyle, formed by the loss of
plate 3", is typically present. One detached operculum has been located and bears
five processes.
DIMENSIONS. Range of observed specimens: shell length 40 (49-6) 62 /*, width
40 (48-4) 58 ^, length of processes 27 (35-6) 48 //,. Number of specimens measured, 11.
OCCURRENCE. H. voigti is rare to fairly common and found spasmodically through-
out the Cenomanian of Fetcham Mill, Compton Bay and Escalles. It was originally
described by Alberti (1961) from the Lower Barremian to ? Lower Aptian of Germany.
The specimens described by Sarjeant (19660) are from the Cenomanian of Fetcham
Mill.
Hystrichodinium dasys sp. nov.
(PI. 10, figs. 8, 9)
DERIVATION OF NAME. Latin, dasys, hairy or shaggy — with reference to the
abundant hair-like processes.
176 CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i
DIAGNOSIS. Shell subspherical, thin-walled, smooth or slightly granular. Tabula-
tion very faintly marked ; cingulum, and more rarely, precingular and postcingular
plate boundaries visible. Processes numerous, short, fine and flexuous, aligned
along plate boundaries, also intratabular. Archaeopyle not observed.
HOLOTYPE. B.M. (N.H.) ¥.51982 (3). Lower Chalk, Bureau de Recherches Geo-
logiques et Minieres Borehole, Escalles, Pas de Calais at 159 metres depth. Upper
Cretaceous (Cenomanian).
DIMENSIONS. Holotype : diameter of central body 58 by 61 p, length of processes
10-14 p. Range: diameter of central body 42 (53-0) 70^, maximum length of
processes 7 (10-6) 14 p. Number of specimens measured, 13.
DESCRIPTION. The shell, being thin-walled, is always found deformed. Only
two specimens possess a faint tabulation. This is best seen in the holotype which
has a clear but lightly defined cingulum and two or three precingular and post-
cingular plate boundaries visible. The numerous processes are hair-like, approxi-
mately 0-2 11 wide for most of their length, probably hollow and terminate with a
point. The most characteristic feature of the processes is their extreme flexibility.
REMARKS. The characteristic processes and the presence of a tabulation differ-
entiate H. dasys sp. nov. very easily from all previously described microplankton
species.
OCCURRENCE. This species is rare to common in all samples from Escalles except
the lower three (E 201, E 207, and E 213). One specimen was recorded from Comp-
ton Bay (sample CB 7) and two from Fetcham Mill (sample FM 520, Turonian).
The species was not recorded from the North American material. The distribution
of H. dasys possibly indicates an environmental difference between Escalles and
Fetcham Mill/Compton Bay, perhaps in the depth of water or distance from land.
IV. REFERENCES
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28 : 93-105, pis. 8, 9.
1961. Zur Kenntnis mesozoischer und alttertiarer Dinoflagellaten und Hystrichos-
phaerideen von Nord- und Mitteldeutschland sowie einigen anderen europaischen Gebieten.
Palaeontographica, Cassel, Stuttgart, 116, A : 1-58, pis. 1-12.
BALTES, N. 1963. Dinoflagellate si Hystrichosphaeride cretacice din Platforma moezica.
Petrol Gaze, Bucuresti, 12 : 581-597, pis. 1-8.
1965. Observatii asupra microflorei cretacice inferiare din zona R. Bacaz. Petrol Gaze,
Bucuresti, 16 : 3-17, pis. 3, 4.
BARROIS, C. 1876. Recherches sur le terrain Cretac6 Superieur de 1'Angleterre et de 1'Irlande.
Mem. Soc. geol. N., Lille, 3, 189-205.
BRISTOW, H. W. 1889. The Geology of the Isle of Wight. Mem. geol. Surv. U.K., London.
BROWN, C. A. 1960. Palynological Techniques, i-vi, 1-188 Baton Rouge, Louisiana.
BROWN, R. W. 1956. Composition of Scientific Words. 882 pp. Reese Press, Baltimore, Md.
CHATTON, E. 1952. Dinoflagelles, in Traite de Zoologie, I, 309-390. Masson, Paris.
CLARKE, R. F. A. & VERDIER, J. P. 1967. An investigation of microplankton assemblages
from the Chalk of the Isle of Wight, England. Verh. K. ned. Akad. Wet., Amsterdam,
24, 3 : 1-96, pis. 1-17.
CENOMANIAN NON-CALCAREOUS MICROPL ANKTON, i 177
CLARKE, R. F. A., DAVEY, R. J., SARJEANT, W. A. S. & VERDIER, J. P. 1968. A Note on the
Nomenclature of some Upper Cretaceous Dinoflagellate Taxa (in press) .
COOKSON, I. C. 1956. Additional microplankton from Australian late Mesozoic and Tertiary
sediments. Aust. J. mar. Freshwat. Res. Melbourne, 7, i : 183-191, pis. i, 2.
— 1965. Cretaceous and Tertiary Microplankton from South-Eastern Australia. Proc. R.
Soc. Viet., Melbourne, 78, i : 85-93, pis. 9-11.
COOKSON, I. C. & EISENACK, A. 1958. Microplankton from Australian and New Guinea Upper
Mesozoic sediments. Proc. R. Soc. Viet., Melbourne, 70, i : 19-79, pis. 1-12.
— igdoa. Microplankton from Australian Cretaceous sediments. Micropaleontology ,
New York, 6, i : 1-18, pis. 1-3.
- 19606. Upper Mesozoic microplankton from Australia and New Guinea. Palaeontology,
London, 2, 2 : 243-261, pis. 37-39.
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PLATE i
Gonyaulacysta exilicristata sp. nov.
Fetcham Mill Borehole (depth, 730 feet).
FIG. i. Dorsal surface of holotype. Slide PF. 3987 (i). x 500.
FIG. 2. Ventral surface. Slide PF. 3987. X 500.
Carpodinium obliquicostatum Cookson & Hughes
FIG. 3. Lower Chalk, Fetcham Mill Borehole (depth, 750 feet). Lateral view. Slide
PF. 3988. x 500.
FIG. 4. Lower Chalk, Escalles Borehole (depth, 159 metres) . Dorsal surface with archaeopyle.
Slide E 195/3. X I25°
Histocysta pa I la sp. nov.
Fetcham Mill Borehole.
FIG 5. Holotype; view of archaeopyle and attached operculum. Slide PF. 3052. x 500.
FIG. 6. Paratype; Antapical view. Slide PF. 3991 (i). X 500.
Gonyaulacysta delicata sp. nov.
FIG. 7. Ventral surface of holotype. V. 51979 (i). X 500.
FIG. 8. Paratype; V. 5 1979(2). X 500.
Gonyaulacysta sp. A. Figured specimen.
Lower Chalk, Fetcham Mill Borehole.
FIG. 9. Ventral surface. Slide PF. 3987 (2). x 500
FIG 10. Medial view. Slide PF. 3987 (2). x 500.
Butt. Br. Mus. nat. Hist. (Geol.) 17, 3
• *
PLATE i
m
4
GEOL. 17, 3.
PLATE 2
Cribroperidiniutn intricatum sp. nov.
FIG. i. Ventral surface of holotype. V. 51980 (i). x 500.
FIG. 2. Lower Colorado, Saskatchewan (depth, 1,023 feet). Dorsal surface with operculum
in situ. ¥.51980. x 500.
FIG. 3. Lower Colorado, Saskatchewan. Detached operculum. Slide Sas 1023/3. X 500.
Microdinium setosum Sarjeant.
FIG. 4. Lower Chalk, Fetcham Mill Borehole (depth, 840 feet). Lateral view. Slide
PF. 3036. x 640.
Microdinium variospinum sp. nov.
FIG. 5. Lower Chalk, Escalles Borehole (depth, 195 metres). Antapical sutural spines
visible. Slide E 195/3. X 640.
FIG. 6. Lateral view of holotype. V. 51981 (i). x 640.
IMicrodinium crinitutn sp. nov.
FIG. 7. Lower Chalk, Fetcham Mill Borehole (depth, 770 feet). Slide FM 770/15. x 640.
FIG. 8. Dorsal surface of holotype Slide PF. 3990 (i). x 640.
Microdinium distinctum sp. nov.
FIG. 9. Ventral surface of holotype. Slide PF. 3989 (i). X 640.
FIG. 10. Dorsal surface of holotype. X 640.
FIG. ii. Lower Chalk, Fetcham Mill Borehole (depth, 790 feet). Ventral surface. Slide
FM 790/16. x 640.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 3
PLATE 2
f
8
v ' -
V
W
10
11 t V
PLATE 3
Ellipsodinium rugulosum Clarke & Verdier
FIG. i. Lateral view of specimen showing operculum partially detached. Slide PF. 3988.
X 1250.
Fromea amphora Cookson & Eisenack
Lower Chalk, Fetcham Mill Borehole.
FIG. 2. Slide FM 770/6 (depth, 770 feet). X 500.
FIG. 3. Specimen with operculum attached (depth, 650 feet). Slide PF. 3041. x 500.
Microdinium veligerum (Deflandre).
FIG. 4. Lower Chalk, Fetcham Mill Borehole (depth, 650 feet). Antapical view to show
crestal cavities. Slide FM 650/5. x 1250.
Apteodinium granulatum Eisenack.
FIG. 5. Lower Chalk, Fetcham Mill Borehole (depth, 690 feet). Ventral surface. Slide
FM 690/12. x 640.
FIG. 6. Lower Chalk, Escalles Borehole (depth, 165 metres). Lateral view. V. 51981.
X 640.
Cassiculosphaeridia reticulata sp. nov.
FIG. 7. Lower Chalk, Fetcham Mill Borehole (depth, 730 feet) . Operculum partially detached.
Slide FM 730/9. X 500.
Chytroeisphaeridia euteiches sp. nov.
FIG. 8. Holotype showing archaeopyle. V. 51982 (2). x 500.
FIG. 9. Lower Chalk, Escalles Borehole (depth, 159 metres). Archaeopyle with detached
operculum. V. 51982. x 500.
Epelidosphaeridia spinosa (Cookson & Hughes)
Lower Chalk, Fetcham Mill Borehole.
FIG. 10. Dorsal surface. Slide PF. 3992, (depth, 770 feet), x 500.
FIG. ii. Ventral surface with sulcus. Slide PF. 3992 (depth, 770 feet). X 500.
FIG. 12. Complete specimen. Slide PF. 3048 (depth, 770 feet), x 500.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 3
PLATE 3
m f
*
i
•A
\
I
8
•x-
-JW
- •••" ':
10
11
12
PLATE 4
Hystrichosphaeridium deanei Davey & Williams
FIG. i. Lower Chalk, Compton Bay. Complete specimen (137 feet above base of Chalk).
Slide CB 19/2. x 500.
Hystrichosphaeridium difficile Manum & Cookson
Upper Colorado, Saskatchewan.
FIG. 2. Complete specimen, archaeopyle partially detached. ¥.51983 (depth, 805 feet).
X 500.
FIG. 6. Detached operculum bearing 4 processes. Slide Sas 805/3 (depth, 805 feet). X 500.
FIG. 7. Detached operculum, lateral view. Slide Sas 805/3 (depth, 805 feet). X 500.
X 500.
Cassiculosphaeridia reticulata sp. nov.
FIG. 3. Holotype; view of archaeopyle. V. 51981 (4). x 500.
Microdinium veligerum (Deflandre).
FIG. 4. Lower Chalk, Fetcham Mill Borehole (depth, 750 feet). Dorsal surface. Slide
PF. 3988. x 640.
Microdinium cf . ornatum Cookson & Eisenack
FIG. 5. Lower Chalk, Escalles Borehole (depth, 189 metres). Dorsal surface. Slide E 189/4.
X 640.
Hystrichosphaeridium radiculatum Davey & Williams
FIG. 8. Lower Chalk, Escalles Borehole (depth, 159 metres). Lateral view showing archaeo-
pyle. V. 51982. x 500.
Hystrichosphaeridium mantelli Davey & Williams.
FIG. 9. Lower Chalk, Escalles Borehole (depth, 153 metres). Lateral view to show archaeo-
pyle. Slide E 153/3. X 500.
Polysphaeridium laminaspinosum Davey & Williams
Lower Chalk, Fetcham Mill Borehole.
FIG. 10. Apical view showing archaeopyle. Slide PF. 3035 (depth, 840 feet), x 500.
FIG. ii. Antapical view. Slide PF. 3035 (depth, 840 feet), x 500.
Bull. Br. Mus. not. Hist. (Geol.) 17, 3
PLATE 4
8
10
11
PLATE 5
Oligosphaeridium anthophorurn (Cookson & Eisenack)
Lower Colorado, Saskatchewan.
FIG. i. Enlargement to show process extremities. Slide Sas 1023/3 (depth, 1023 feet).
X 975-
FIG. 2. Detached operculum. Slide Sas 1023/1 (depth, 1023 feet). X 500.
FIG. 3. V. 51980 (boring depth, 1023 feet), x 500.
Oligosphaeridium prolixispinosum Davey & Williams
FIG. 4. Lower Chalk, Compton Bay (116 feet above the base of the Chalk). Specimen
showing " bald " cingular region. Slide CB I7/C. X 500.
Hystrichosphaeridium tubiferum (Ehrenberg)
Lower Chalk, Fetcham Mill Borehole.
FIG. 5. Detached operculum. Slide FM 690/12 (depth, 690 feet). X 500.
FIG. 8. Slide PF. 3987 (depth, 730 feet), x 500.
Oligosphaeridium complex (White)
FIG. 6. Lower Chalk, Escalles Borehole (depth, 159 metres). Detached operculum composed
of 4 plates. V. 51982. x 500.
FIG. 7. Lower Chalk, Fetcham Mill Borehole (depth, 750 feet). Specimen illustrating the 6
precingular processes. Slide FM 750/13. x 500.
Hystrichosphaeridium bowerbanki Davey & Williams
FIG. 9. Lower Chalk, Compton Bay (116 feet above the base of the Chalk). Lateral view,
archaeopyle to the north. Slide CB I7/C. x 500.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 3
PLATE 5
.
'/
5
GEOL. 17, 3.
PLATE 6
Oligosphaeridiutn reniforme (Tasch)
FIG. i. Lower Colorado, Saskatchewan (depth, 1023 feet). Slide Sas 1023/1. x 500.
Tanyosphaeridiutn variecalamum Davey & Williams
FIG 2 Holotype. Slide PF 3035 (2) x 500
FIG 5 Lower Chalk, Compton Bay (28 feet above base of Chalk). Precingular, cingular and
postcingular processes aligned into three series. Slide CB 5/C x 500
Litosphaeridium siphoniphorum (Cookson & Eisenack)
Lower Chalk, Fetcham Mill Borehole.
FIG. 3. Lateral view showing precingular and postcingular processes, and antapical process.
Slide PF. 3987 (depth, 730 feet), x 500.
FIG. 4. Detached operculum. Slide FM 690/14 (depth, 690 feet). X 975.
Callaiosphaeridium asymmetricum (Deflandre & Courteville) .
FIG. 6. Upper Greensand, Fetcham Mill Borehole (depth, 886 feet). Lateral view illustrating
epitractal archaeopyle. Slide FM 886/2. x 500.
Cleistosphaeridiutn polypes (Cookson & Eisenack)
FIG. 7. Upper Woodbine Formation. Enlargement to show process extremities. Slide
T5/3- X 975-
FIG. 8. Lower Chalk, Compton Bay (151 feet above base of Chalk). Specimen possessing
apical archaeopyle. Slide CB 2i/C. x 500.
Cleistosphaeridiutn polypes var. clavulum nov.
FIG. 9. Type. Slide PF. 3995 (i) x 500.
FIG. 10. Lower Chalk, Fetcham Mill Borehole (depth, 840 feet). Enlargement to show
process extremities. Slide PF. 3035. x 975.
fCleistosphaeridium aciculare sp. nov.
FIG. ii. Upper Colorado, Saskatchewan (depth, 835 feet). Specimen with numerous
fine processes. V. 51988. x 500 (phase contrast).
FIG. 12. Holotype. V. 51979 (3). X 500.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 3
PLATE 6
8
12
, *t
10
PLATE 7
Exochosphaeridium striolatutn var. truncatum nov.
FIG. i. Lower Chalk, Fetcham Mill Borehole (depth, 750 feet). Apical view showing two
partially detached precingular plates to the north. Slide PF. 3988. x 500.
FIG. 2. Type; precingular archaeopyle clearly shown to be formed by the removal of two
plates. V. 51982 (i). x 500.
FIG. 3. Lower Chalk, Escalles Borehole (depth, 165 metres). Detached operculum composed
of two precingular plates. ¥.51981. x 500.
fCleistospaeridium flexuosum Davey et al.
FIG. 4. Lower Chalk, Fetcham Mill Borehole (depth, 690 feet). Slide PF. 3030. x 500
(phase contrast).
Exochosphaeridium phragmites Davey et al.
FIG. 5. Lower Chalk, Escalles Borehole (depth, 165 metres). Apical process with precingular
archaeopyle to the north-west. V. 51981. x 500.
Cleistosphaeridium huguonioti var. pertusutn nov.
FIG. 6. Lower Chalk, Fetcham Mill Borehole (depth, 690 feet). Enlargement showing
vacuolated processes with small distal bifurcations. Slide FM 690/14. x 975.
FIG. 7. Type. Slide PF. 3040 (2). x 500.
FIG. 9. Lower Chalk, Escalles Borehole (depth, 159 metres). Specimen possessing apical
archaeopyle. Slide E 159/4. X 500.
Cleistosphaeridium heteracanthum (Deflandre & Cookson)
FIG. 8. Lower Chalk, Compton Bay (137 feet above base of Chalk). Slide CB 19/2. x 500.
Cleistosphaeridium huguonioti (Valensi)
FIG. 10. Lower Chalk, Fetcham Mill Borehole (depth, 730 feet). Detached operculum.
Slide FM 730/19. x 975.
? Cleistosphaeridium parvum sp. nov.
FIG. ii. Holotype. V. 51981 (3). x 975.
FIG. 12. Lower Chalk, Escalles Borehole (depth, 159 metres). Cingular region devoid of
processes clearly visible. Slide E 159/1. X 975.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 3
PLATE 7
/
k
*
' * 7
PLATE 8
Cleistosphaeridiutn armatum (Deflandre)
FIG. i. Lower Chalk, Escalles Borehole (depth, 189 metres). View of apical archaeopyle.
Slide E 189/4. x 975-
FIG. 2. Lower Chalk, Compton Bay (116 feet above base of Chalk). Enlargement to show
process extremities. Slide CB iy/C. x 975.
FIG. 12. Lower Chalk, Compton Bay (i 16 feet above base of Chalk). Slide CB I7/C. x 500.
Cyclonephelium eisenacki sp. nov.
FIG. 3. Lower Colorado, Saskatchewan (depth, 1,023 feet). Specimen with archaeopyle
developed. ¥.51980. X 500.
FIG. 4. Holotype. V. 51980 (2). x 500.
Prolixosphaeridium conulurn sp. nov.
FIG. 5. Holotype; complete specimen. V. 51981 (5). x 500.
FIG. 6. Lower Chalk, Compton Bay (59 feet above base of Chalk). Apical archaeopyle to
the north. Slide CB 9/2. X 500.
Cleistosphaeridium multifurcatum (Deflandre)
FIG. 7. Lower Chalk, Escalles Borehole (depth, 207 metres). View of apical archaeopyle.
Slide E 207/5. X 500.
FIG. 10. Lower Chalk, Compton Bay (15 feet above base of Chalk). Complete specimen.
V. 51986. x 500.
Coronifera oceanica Cookson & Eisenack
FIG. 8. Lower Chalk, Fetcham Mill Borehole (depth, 730 feet). Specimen possessing an
apical archaeopyle and an antapical process. Slide FM 730/13. x 500.
FIG. ii. Lower Chalk, Escalles Borehole (depth, 189 metres). Complete specimen. Slide
E 189/4. x 500.
Surculosphaeridiutn longifurcatum (Firtion)
FIG. 9. Lower Chalk, Fetcham Mill Borehole (depth, 730 feet). Lateral view to show apical
archaeopyle, deeply furcate precingular processes and completely subdivided cingular processes.
Slide PF. 3987. X 500.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 3
PLATE 8
•** ••••
10
12
PLATE 9
Cyclonephelium paucispinum sp. nov.
FIG. i. Holotype. V. 51981 (2). x 500.
FIG. 2. Lower Chalk, Compton Bay (15 feet above base of Chalk). Complete specimen with
operculum partially detached. V. 51986. x 500.
Cyclonephelium vannophorum sp. nov.
FIG. 3. Enlargement of holotype to show apical archaeopyle, and the shape of the processes.
V. 51986 (i). X 975-
Cyclonephelium eisenacki sp. nov.
FIG. 4. Lower Colorado, Saskatchewan (depth, 1,023 feet). Complete specimen. Slide
Sas 1023/3. X 500.
Hystrichokolpoma ferox (Deflandre)
Lower Chalk, Fetcham Mill Borehole (depth, 840 feet). Slide FM 840/11.
FIG. 5. Lateral view (bottom of specimen by transparency) showing precingular, cingular
and postcingular processes ; large antapical process to the south and fine sulcal processes to the
west, x 500.
FIG. 6. Medial section, x 500.
FIG. 7. Lateral view (top of specimen) showing precingular, cingular and postcingular
processes, x 500.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 3
PLATE 9
3
v
PLATE 10
Hystrichosphaera ramosa var. ramosa (Ehr.)
Lower Chalk, Fetcham Mill Borehole (depth, 750 feet).
FIG. i. Ventral surface. Slide PF. 3988. x 500.
FIG. 2. Dorsal surface with precingular archaeopyle. Slide PF. 3988. x 500.
FIG. 5. Detached operculum (boring depth, 770 feet). Slide FM 770/15. x 500.
Hystrichosphaera ramosa var. multibrevis Davey & Williams
Lower Chalk, Fetcham Mill Borehole.
FIG. 3. Slide PF. 3988 (depth, 750 feet), x 500.
FIG. 4. Slide PF. 3988 (depth, 750 feet), x 500.
Hystrichodiniutn voigti (Alberti)
Lower Chalk, Escalles Borehole.
FIG. 6. Archaeopyle to the north-east. V. 51982 (depth, 159 metres).
FIG. 10. Detached operculum bearing 4 processes. Slide E 165/1 (depth, 165 metres).
X 500.
Achomosphaera ramulifera (Deflandre)
FIG. 7. Lower Chalk, Escalles Borehole (depth, 159 metres). Specimen showing precingular
archaeophyle and apical process. V. 51982. x 500.
Hystrichodinium dasys sp. nov.
FIG. 8. Holotype illustrating cingulum. V. 51982 (3). x 500.
FIG. 9. Middle Chalk, Fetcham Mill Borehole (depth, 520 feet). Specimen with unusually
stout processes proximally. Slide FM 520/7. x 500.
Adnatosphaeridium chonetum (Cookson & Eisenack)
FIG. ii. Lower Chalk, Escalles Borehole (depth, 165 metres). Complete specimen, V. 51981.
X 500.
FIG. 12. Lower Chalk, Fetcham Mill Borehole (depth, 730 feet). Apical archaeopyle present.
Slide PF. 3987. x 500.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 3
PLATE 10
8
10
--•
12
PLATE ii
Trichodinium castaneum (Deflandre).
Lower Chalk, Escalles Borehole.
FIG. i. Lateral view showing precingular archaeopyle and cingulum, V. 51989 (depth,
195 metres), x 500.
FIG. 2. Dorsal view showing small apical horn. Slide E 159/2 (depth, 159 metres), x 500.
FIG. 3. Complete specimen possessing a cingulum and possibly a sulcus, Slide E 189/4 (depth,
189 metres), x 500.
Exochosphaeridium pseudohystrichodinium (Deflandre)
Lower Chalk, Escalles Borehole.
FIG. 4. Complete specimen with operculum partially detached, V. 51982 (depth, 159 metres).
X 500.
FIG. 5. Complete specimen illustrating pitted surface, V. 51981 (depth, 165 metres), x 500.
Cyclonephelium distinctum Deflandre & Cookson
FIG. 6. Lower Chalk, Escalles Borehole (depth, 159 metres). Specimen with unusually
long processes, V. 51982. x 500.
FIG. 7. Lower Chalk, Escalles Borehole (depth, 159 metres). Complete specimen; archaeo-
pyle in the act of developing. V. 51989. x 500.
FIG. 8. Lower Chalk, Compton Bay (15 feet above base of Chalk). Typical specimen with
archaeopyle developed. V. 51986. X 500.
FIG. 10. Lower Chalk, Escalles Borehole (depth, 165 metres). Detached operculum.
V. 51981. x 500.
Cyclonephelium membraniphorum Cookson & Eisenack
FIG. 9. Lower Chalk, Escalles Borehole (depth, 153 metres). Apical archaeopyle well
illustrated. Slide E 153/3. X 500.
Cyclonephelium vannophorum sp. nov.
FIG. ii. Holotype with operculum partially detached. V. 51986 (i). x 500.
FIG. 12. Lower Chalk, Compton Bay (15 feet above base of Chalk). Specimen with apical
archaeopyle developed. V. 51986. x 500.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 3
PLATE ii
\ \
•
w?
10
PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING
A REDESCRIPTION OF
W. CARRUTHERS' TYPE
GRAPTOLITES
ISLES STRACHAN
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 4
LONDON: 1969
A REDESCRIPTION OF W. CARRUTHERS'
TYPE GRAPTOLITES
BY
ISLES STRACHAN
Pp. 181-206; 5 Plates; 8 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 4
LONDON : 1969
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 17, No. 4 of the Geological
(Palaeontological) series. The abbreviated titles of
periodicals cited followthose of the World List of
Scientific Periodicals.
World List abbreviation:
Bull. Br. Mus. nat. Hist. (Geol.).
Trustees of the British Museum (Natural History) 1969
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 17 January, 1969 Price £i 45.
A REDESCRIPTION OF W. CARRUTHERS'
TYPE GRAPTOLITES
By ISLES STRACHAN
CONTENTS
Page
I. INTRODUCTION ......... 183
II. DESCRIPTIONS ......... 185
Leptograptus capillaris . . . . . . . 185
Pleurograptus linearis . . . . . . . 186
Dicellograptus elegans . . . . . . . 187
Dicellograptus moffatensis . . . . . . . 189
Dicranograptus clingani . . . . . . . 190
Climacograptus minimus ....... 191
Climacograptus minutus . . . . . . . 193
Cvyptograptus tricornis . . . . . . . 194
Monograpius clingani . . . . . . . 195
Monograptus intermedius . . . . . . . 196
Diver sograptus? capillaris . . . . . . . 198
Rastrites maximus ........ 200
Cyvtograptus murchisoni ....... 202
" Dendrograptus " lentus . . . . . . . 204
III. REFERENCES .......... 204
SYNOPSIS
The fourteen species of graptolites described and figured by William Carruthers, mainly in
his paper of 1868, are redescribed from the type material where possible. Two neotypes are
proposed. Some of the species appear to be unsatisfactory by modern standards, but it is
hoped that refiguring of the types will assist future work.
I. INTRODUCTION
IN their monograph of British Graptolites (1901-18) Elles & Wood described and
figured most of the previously known species as well as numerous new forms. In
many cases, however, they did not use the original type material either because it
was not available or because later collections had provided other, sometimes better,
material. In the fifty years which have elapsed since then, discrimination of grapto-
lite species has become keener, particularly by workers overseas who have frequently,
in their subdividing, formally designated type specimens without consideration of
the availability or value of the early material. It seems appropriate that the species
described in the nineteenth century should be re-examined so that these workers
should be able to recognize the old species accurately with the aid of modern
descriptions and illustrations. The work could be tackled systematically by genus
and species but this involves consideration of generic and specific limits in deciding
GEOL. IJ, 4 15
184 W. CARRUTHERS' TYPE GRAPTOLITES
the scope of a paper. It is easier to treat the species for redescription by discussing
one author at a time and I hope to undertake further papers of a similar scope to this.
William Carruthers (1830-1922) was born at Moffat, Dumfriesshire, and early
developed an interest in the local geology, particularly in the graptolites for which
the area had then recently become famous through the researches of Harkness and
Salter. His own publications on graptolites are comparatively few in number but
it is clear that by 1867 he was regarded as an expert on the group, since he contributed
the article on graptolites to the fourth edition of Murchison's " Siluria ". In the
following year he described more fully the new species erected in that memoir.
He strongly disagreed with Nicholson on the interpretation of a number of graptolite
features and they carried on an acrimonious correspondence in the pages of the
Geological Magazine during 1867. At the same time he was also describing fossil
plants, to which he later devoted all his research. In 1870-72 he was assisting the
young amateur John Hopkinson with his descriptions of new species of graptolites
from the Moffat Shales but they both appeared to realize that the publication in
1876 of Lap worth's paper on " Scottish Monograptidae " marked a new and more
critical approach to graptolite systematics and stratigraphy and neither of them
published further papers on the subject. It is, however, clear from Lapworth's
later papers that he maintained a friendly association with Hopkinson and Carruthers
and the latter's manuscript notes on graptolites are in the Lapworth Library,
Birmingham University. These notes contain no indication of how or when Lap-
worth acquired them but there are no references to papers after 1872 and it is possible
that Carruthers gave them to Lapworth before taking up more administrative
duties at the British Museum (Natural History) in 1880 and confining his research
to palaeobotany.
The new species described by Carruthers are found in two main papers, in 1858
and 1868. The first of these was reprinted the following year in a more accessible
journal and, as already mentioned, the 1868 paper described properly the species
which were only illustrated in the appendix to " Siluria ". Carruthers had some
harsh remarks to make about Nicholson's drawings of graptolites and, on the whole,
his own are generally sufficiently accurate to allow recognition of the type specimens.
Most of these were catalogued at once in the British Museum (Natural History)
collections where they can be traced. Almost all his species can be recognized
fairly easily. The type of Rastrites maximus is the only one which has not been
found and a neotype is proposed for that species.
It is a pleasure to acknowledge the ready co-operation which I have had from those
at the British Museum (Natural History) who have had charge of the graptolite
collections and to thank their photographers who have provided most of the illus-
trations for the plates. I must also express my thanks to the Geological Survey &
Museum and to the Royal Scottish Museum for the loan of specimens over a period
which has been much longer than I had at first intended.
Specimens with Q numbers are in the British Museum (Natural History); with
GSM, in the Institute of Geological Sciences, London; with BU, in the Geology
Department, University of Birmingham; and with SM, in the Sedgwick Museum,
Cambridge.
W. CARRUTHERS' TYPE GRAPTOLITES 185
II. DESCRIPTIONS
Leptograptus capillaris (Carruthers)
PL i
1868 Cladograpsus capillaris Carruthers: 130, pi. 5, figs. 7, ya.
1876 Leptograptus capillaris (Carr.) Lapworth : 9, pi. 3, fig. 72.
1903 Leptograptus capillaris (Carr.); Elles and Wood : 112, pi. XV, figs. 4a-d.
?I954 Leptograptus capillaris (Carr.); Sherrard 195, pi. n, fig. 8.
Original description. " Extremely slender polypary, with remote branches and
very minute hydrothecae; about twenty-four in an inch. It is not so abundant as
C. linearis, and is easily distinguished by its capillary appearance. It is probably
the same species as that figured and described by Emmons in his American Geology,
vol. i, p. 109, pi. i, fig. 7, under the name of Nemagrapsus capillaris. Loc. Moffat ".
Comments on usage. There appears to have been little confusion over this species.
Although he quoted Emmons' species, Carruthers clearly intended his own form
to rank as a new species and the American form is now referred to Thamnograptus.
The original description refers to " remote branches " which are not shown by the
type specimens or original figure but one illustration of Cladograpsus linearis of the
previous year (Carruthers 1867, pi. 2, fig. I7a) shows the characteristic curved
stipes with secondary branches. The original of this has not however been
recognized.
TYPE MATERIAL. The syntypes, British Museum (Natural History) Q-30, were
refigured by Elles & Wood. As lectotype I select the large rhabdosome on the
upper right of Carruthers' original figure (see PI. i, fig. 2). The precise locality and
horizon are not known but Elles & Wood give it as Hartfell Spa?, Hartfell Shales.
DIAGNOSIS. Very slender Leptograptus with markedly curved stipes, maximum
breadth 0-5 mm. thecae eight to ten per cm.
REVISED DESCRIPTION. The species normally occurs crowded on the slabs so that
details of the stipes are not readily seen. The sicula is inconspicuous but the stipes
appear to grow initially horizontally or even somewhat downward before curving
gracefully upwards and inwards, forming loops on occasion. The stipes widen from
an initial breadth of 0-2 mm. to a maximum of 0-5 mm. over a distance of at least
4 or 5 cm. and appear to be twisted so that the thecae can appear on either side of
the curve. The sicula is 2 mm. long and about 0-2 mm. wide but details of the origin
of the stipes are not known. The thecae appear to be simple leptograptid tubes
but the preservation of the distal parts of the stipes is such as to make measurements
of their length and overlap impossible.
GEOLOGICAL HORIZON. Lapworth (1878) recorded the species only from the
zone of Pleurograptus linearis but Elles & Wood (1903) also recorded it from the zone
of Dicranograptus clingani. The type slab shows no associates, nor do most of the
specimens in the Lapworth Collection. Lithologically, however, the slabs agree
with the upper part of the Lower Hartfell Shales.
DISTRIBUTION. This species is known from only a few localities in the South of
Scotland and is not recorded from beds of a similar age in South Wales. Ruedemann
i86 W. CARRUTHERS' TYPE GRAPTOLITES
and Decker recorded it from the Viola Limestone in Oklahoma but gave no figures
(Ruedemann 1947). Sherrard (1954) recorded it from New South Wales but her
figure shows stipes which appear to be too broad and not curved enough. They
may be one of the forms of Leptograptus flaccidus which occurs about the same hori-
zon, e.g. L.f. arcuatus Elles & Wood. Thomas (1960) illustrated a form which is
clearly not L. capillaris as the stipes are too straight, and this then casts doubt on
the range of the species as given by Thomas, which goes up to the Ashgill Series.
Pleurograptus linearis (Carruthers)
PL 2 ; Figs. la-c
1858 Cladograpsus linearis Carruthers : 467, fig. i.
1859 Cladograpsus linearis Carruthers : 24, fig. i.
non 1867 Cladograpsus linearis Carruthers; Carruthers : 369, pi. 2, fig. 17.
?i8&7a Cladograpsus linearis Carruthers; Carruthers : 540, fig. 8.
1867 Pleurograpsus linearis (Carruthers) Nicholson : 257, pi. n, figs. 1-5.
1868 Cladograpsus linearis Carruthers ; Carruthers : 129.
1876 Pleurograptus linearis (Carruthers) ; Lapworth, pi. 3, fig. 69.
1904 Pleurograptus linearis (Carruthers); Elles & Wood : 119, pi. 16, fig. 7, pi. 17, fig. i.
ORIGINAL DESCRIPTION. " From a short and very slender base the zoophyte
divides into two stems, each supporting the cells on their upper sides. Branches
are given off at irregular intervals from these principal stems. The length of the
polypidom is very great; one specimen I have been able to trace for nearly three
feet . . . The polypidom at its origin, near to the slender base, is very narrow,
being little more than a fine line; as it increases in length it increases in breadth,
until it is fully two-fifths of a line broad. The cells are very remote from each
other, and are, at first sight, from the slight indentation they make in the stem,
scarcely perceptible, giving the Graptolite the appearance as if it were a clear line.
The mouth of the cell is straight and at right angles to the axis ; it makes an inden-
tation equal to about one-sixth of the breadth of the polypidom. The number of
cells in an inch is about eighteen. Type locality Hartfell."
COMMENTS ON USAGE. Carruthers' later figures (? 1867, i867a) are different from his
original but there has been no difficulty in the recognition of his species. Carruthers
accepted Nicholson's specimen of Pleurograpsus linearis although he was very critical
of the detail of the figures. Nicholson's specimen was refigured by Elles & Wood
who also figured a variety simplex in which the branches are much more widely
spaced.
TYPE MATERIAL. Carruthers' original specimen is in the British Museum (Natural
History), (^.848, and the counterpart is amongst material presented to the Royal
Scottish Museum, Edinburgh, by Carruthers in 1858 (RSM 1858 . 10 .4). Nicholson's
specimen, Q . 27, was said by Elles & Wood to be the type specimen but it is clearly
not. The lower of the two large specimens on the type slab has four lateral branches
and is recognizable as the specimen figured in 1858 (see PL 2, figs, i, 2).
REVISED DESCRIPTION. Rhabdosome consisting of two or three main stipes
arising from the sicula with secondary branching on one or both sides. Stipes up
W. CARRUTHERS' TYPE GRAPTOLITES
187
to I mm. broad; thecae simple leptograptid, about eight per cm., with apertural
excavations occupying about one-third of the rhabdosome breadth. The sicula
is not seen on any of the figured material and is probably associated in older rhabdo-
somes with one of the branches, a feature also seen in species of Leptograptus where
a third branch may be developed. Young specimens on the type slab, however,
suggest that the sicula is about 2 mm. long. The stipes are initially very slender
but show some secondary thickening in older specimens where the thecae on the
main stipes are difficult to distinguish. There is considerable variation in the
branching, Nicholson's specimen (PL 2, fig. 3) being much stiffer than Carruthers'
or the other specimen figured by Elles & Wood in which the tertiary branched stipes
are aligned in flowing curves. This suggests that the variation in attitude of the
stipes is simply a depositional effect.
fl
^
FIG. i. Pleurograptus linearis Carr. a. Enlargement of proximal part of main stipe showing
2 lateral branches and possible central branch from the sicula. Counterpart of type slab,
R.S.M. 1858.10.4. The 2 parallel distal parts of branches are the left hand pair from the
holotype (PI. 2, fig. i). b, c. Young specimens showing sicula, type slab, Q.8^8. All X5-
DISTRIBUTION. P. linearis appears to be confined to the one horizon in the south
of Scotland where, however, it is quite common if all the fragments showing this
type of branching really belong to it. Ruedemann (1908; 1947) figured a single
specimen from the Utica Shale of New York State which shows similar branching
at rather more distant intervals, approximating in that respect to var. simplex
which is also recorded by Thomas (1960) from Australia.
Dicellograptus elegans (Carruthers)
PI. 3, fig. i. ; Figs 2a, b.
Didymograpsus elegans Carruthers : 369, pi. 2, fig. i6a.
Didymograpsus elegans Carruthers; Carruthers : 129, pi. 5, figs. 8a, ?d.
Dicellograpsus elegans (Carruthers) Hopkinson : 24, pi. i, fig. 3.
Dicellograptus elegans (Carruthers) ; Lapworth, pi. 4, fig. 87.
Dicellograptus elegans (Carruthers)
Dicellograptus elegans (Carruthers)
1867
1868
1871
1876
1877
1904
?I947
1954
ORIGINAL DESCRIPTION. " Branches of the polypary divaricating at various
angles, and with a slight curve within a short distance of the proximal origin of the
Dicellograptus elegans (Carruthers)
Dicellograptus elegans (Carruthers) ; Sherrard, pi. 10, fig. 6.
Lapworth : 141, pi. 7, fig. 8.
Elles & Wood : 159, pi. 23, figs. 2a-e.
Ruedemann : 380, pi. 63, fig. i.
i88
W. CARRUTHERS' TYPE GRAPTOLITES
polypary. The hydrothecae are rounded at the apex, and free throughout a con-
siderable portion of their length and the intervening spaces are rounded at the base ;
about twenty-two cells in an inch. The initial process is obvious in young specimens,
but I have not been able to detect it in old individuals ; the outer apex of the angle
ornamented with three short strong spines. Loc. Moffat ".
COMMENTS ON USAGE. Since Elles & Wood refigured the type specimen there
has been no difficulty with this species. The originals of Carruthers' figures 8b
and 8c (BM(NH) Q-54) were recognized by Hopkinson (1871) as being a distinct
species (D. morrisii Hopkinson) and not merely young stages of elegans as Carruthers
thought.
TYPE MATERIAL. The holotype, Q.850 (PL 3, fig. i), is from the Hartfell Shales,
Dobbs Linn. The species appears to be commonest in the P. linearis Zone but the
associates on the type slab (Dicellograptus pumilus and Climacograptus spp.) give
no direct confirmation of horizon.
DIAGNOSIS. Dicellograptus with markedly introverted thecae and with distinct
sigmoid curvature of stipes near proximal end, so that the stipes curve first upwards,
then outwards, then upwards again.
FIG. 2. Dicellograptus spp. a. D. elegans Carr. Detail of thecal aperture in proximal part
of stipe. Q.850. x 30. b. D. elegans Carr. Distal thecae. (3.850. X5- c. D. moffatensis
Carr. Distal thecae. (3.843. X5-
REVISED DESCRIPTION. The proximal double curvature is quite distinctive and
the stipes have an almost uniform width of i mm. throughout their length. The
first two thecae have prominent apertural spines which, with the virgella, give the
three-spined proximal end noted by Carruthers. The thecae number eight to ten
per cm. and are of the strongly introverted type with marked ventral curvature
(group IV of Elles & Wood, which however are not introtorted, see Bulman 1944 : 37).
GEOLOGICAL HORIZON. Lapworth (1878) and Elles & Wood (1904) recorded the
species only from the P. linearis zone, but in the summary range chart at the end
of the Monograph, Elles & Wood also recorded it as common in the underlying zone
of D. clingani. Elles (1925) also recorded it from both zones so its precise range
must await a revision of the Hartfell Shales. Foreign records are surprisingly
scanty but it seems to occur in Australia at the same horizon.
W. CARRUTHERS' TYPE GRAPTOLITES 189
Dicellograptus moffatensis (Carruthers)
PL 3, figs. 5, 6; Fig. 2c
1858 Didymograpsus Moffatensis Carruthers : 469, fig. 3.
1859 Didymograpsus Moffatensis Carruthers; Carruthers
1868 Didymograpsus Moffatensis Carruthers; Carruthers
26, fig. 3.
129.
25, pi. i, fig. 4.
1871 Dicellograpsus Moffatensis (Carruthers) Hopkinson
1875 Dicellograptus moffatensis (Carruthers); Hopkinson & Lapworth : 654, pi. 34, fig. 5a.
1877 Dicellograptus Moffatensis (Carruthers); Lapworth : 141, pi. 7, fig. 9.
1904 Dicellograptus moffatensis (Carruthers); Elles & Wood : 157, pi. 23, figs. la-f.
ORIGINAL DESCRIPTION. ' The base terminates in three distinct spinous pro-
cesses. The zoophyte bifurcates from the base. The general appearance is like
the figure; or occasionally the lines form an acute angle for about a quarter of an
inch, then suddenly expand in slight curves, almost at right angles, for a short dis-
tance, when they again recur to their original direction. The branches are united
for about a quarter of a line by a slight web, which in some specimens is terminated
in a fine process of short length, taking the direction of a line bisecting the angle.
The cells are arranged in the outer margins; they are very remote, and penetrate
the polypidom to scarcely one-fourth of its breadth; they form slight openings on
the margin of the polypidom, first entering at a right angle, and then suddenly
turning downwards. These openings are lengthened ovate pouches, answering
exactly in shape and size to the cell-serratures of the margin. The number of cells
in an inch is about twenty. The breadth of the polypidom is about two-thirds of a
line. Type locality Hartf ell ."
COMMENTS ON USAGE. In 1868, Carruthers noted D. divaricatus (Hall) and
D. anceps (Nicholson) as synonyms of his own species but this is merely a reflection of
the confused state of graptolite systematics at the time. Elles & Wood refigured
Carruthers' type but the species does not seem to have been widely recognized.
TYPE MATERIAL. This is one of the few species of which Elles & Wood made any
discussion of type specimens. They considered that Carruthers' specimen was not
a satisfactory type and that a specimen from the Lapworth Collection should be
taken as the type. Carruthers' original figure is admittedly poor (PL 3, fig. 5) but
the type slab shows three specimens which agree fairly well with it and show the
essential characters of the species. The abrupt widening of the stipes on Lap worth's
specimen, which Elles & Wood wanted as a character of the species, appears to be
the result of slight shearing.
It is impossible to decide which of the three specimens was the original of
Carruthers' figure so the specimen (Q . 843) figured by Elles & Wood is here selected as
lectotype (see PL 3, fig. 6). It is from the Hartf ell Shales of Hartf eU and is associa-
ted on the slab with Orthograptus cf . whitfieldi. This probably indicates a low horizon
in the Hartf ell Shales. Lapworth (1878) records it no higher than the zone of
Climacograptus wilsoni.
REVISED DESCRIPTION. Stipes sub-parallel initially, then diverging making an
angle of about 45°, sometimes later converging. Stipes widening from an initial
breadth of 0-4 mm. to a maximum of about 1-5 mm. Thecae eight to ten per cm.,
igo W. CARRUTHERS' TYPE GRAPTOLITES
with markedly curved ventral walls and introverted apertures. The proximal end
usually shows a membrane in the axil of the stipes, obscuring the sicula. The
virgella and first thecal spines variably developed. The thecae are poorly preserved
but distally appear to be very similar to those of D. elegans (Fig. 2c).
It is unfortunate that both Carruthers and Hopkinson included D. divaricatus
(Hall) in their synonymy of this species as it makes it difficult to establish what they
regarded as the diagnostic features of the species. The general shape of the rhab-
dosome appears to be the most characteristic feature, particularly the narrow axil
and later divergence. In this respect the specimen figured by Elles & Wood in
their Monograph (pi. 23, fig. ic) is not at all typical but its only associates are the
pair of specimens figured on the same plate (pi. 23 fig. ib) which have the typical
shape. As the rhabdosomes were flexible in life to some extent, this poses the prob-
lem of how much reliance should be placed on rhabdosome shape as a specific
character in Dicellograptus and other forms with long slender stipes. D. moffatensis
can be readily separated from other British Dicellograptus by size and shape of rhab-
dosome, D. morrisii Hopkinson being the most similar but with a more open axil.
D. moffatensis var. alabamensis Ruedemann 1908 is not related at all and is clearly
a Dicranograptus, close to D. brevicaulis Elles & Wood 1904.
DISTRIBUTION. Elles & Wood recorded the species widely throughout the British
Isles but it seems doubtful now if it occurs in the Lake District. The specimen
figured by Hopkinson & Lapworth from Llanvirn (SM . Ai740o) is a poorly-preserved,
bent dichograptid (O. M. B. Bulman, personal communication). The specimen
from Abereiddy Bay refigured by Elles & Wood may well come from the locality
referred to the " Dicranograptus Shales " (Cox, 1915 : 304) and not from the
D. murchisoni Shales for which the area is best known. If this is so, the species ranges
from PLlandeilo Series to Caradoc Series (wilsoni Zone). It has been recorded from
Australia but Thomas (1960) does not list it in his range chart and so presumably
does not consider it to be present. Linnarsson recorded it from Scania but Hadding
(1913) transferred this form to his new species D. vagus.
Dicranograptus clingani Carruthers
PI. 3, figs. 2-4; Fig. 3a
1868 Dicranograptus Clingani Carruthers : 132, pi. 5, figs. 6a-c.
1870 Dicranograptus Clingani Carruthers; Hopkinson : 358, pi. 16, figs. 4a-c.
1876 Dicranograptus Clingani Carruthers; Lapworth, pi. 3, fig. 76.
1877 Dicranograptus Clingani Carruthers; Lapworth : 141, pi. 6, fig. 43.
1904 Dicranograptus Clingani Carruthers; Elles & Wood : 165, pi. 24, figs. ia-1.
1915 Dicranograptus Clingani Carruthers; Hadding : 22, pi. 3, figs. 1-8.
ORIGINAL DESCRIPTION. " Polypary with a short diprionidian portion, the proxi-
mal end furnished with three very delicate spines; hydrothecae forming a slight
serration along the margin; twenty-one cells in the inch. Loc. Moffat."
COMMENTS ON USAGE. Elles & Wood put this species in a group by itself on the
basis of the thecal characters — approximately straight ventral walls and horizontal
W. CARRUTHERS' TYPE GRAPTOLITES 191
apertures. These characters are clearly seen in Hopkinson's figures and serve to
differentiate the species from other forms with a short biserial portion.
TYPE MATERIAL. Elles & Wood refigured as " type specimen " the original of
Carruthers' fig. 6a and this can be taken as a designation of a lectotype. The
specimen, Q-55 (PI. 3, figs. 3, 4), is from the Hartfell Shales, Hartfell Spa.
Carruthers' fig. 6b, Q.842 (PL 3, fig. 2), is also from this locality.
REVISED DESCRIPTION. Dicranograptus with short biserial portion consisting of
three or four pairs of thecae, and short, straight uniserial stipes diverging at about
40°. The biserial portion has a uniform breadth of about i mm. and the branches
are similarly uniform at about 0-8 mm. The virgella is usually prominent as a
short spine and the first two thecae may have sub-apertural spines. The ventral
walls of the later thecae are straight and the apertures are horizontal in excavations
which occupy about one-third of the breadth. The uniserial stipes appear to be
rarely more than 2 cm. long although Hadding figures a specimen with stipes nearly
4 cm. long. The distal thecae number eight to ten per cm. but the stipes are usually
twisted so that the thecae are in scalariform view (Fig. 3a) and it is difficult to decide
the precise thecal shape. The apertural excavations are clearly marked by lists and
it is probable that the straight ventral walls of the proximal thecae continue in the
distal ones.
FIG. 3. a. Dicranograptus clingani Carr. Proximal end of paratype. (3.842. X5- The
right hand side of the specimen is poorly preserved, b. Climacograptus minutus Carr. Lectotype.
Q8o. X5- c. Cryptograptus tricornis Carr. Proximal end of lectotype showing basal spines.
X5-
DISTRIBUTION. The species is common in the Lower Hartfell Shales of the Moffat
area and is also found at Conway. Elles & Wood recorded it from equivalent beds
in Ireland. It is found in Scandinavia but has not been recorded from North
America. Thomas (1960) gives records but omits it from his range chart, so casting
doubt on the records. The type specimen has no associates but Elles & Wood
recorded it only from its own zonal association.
Climacograptus minimus (Carruthers)
PI. 4, fig. 3; Fig. 4c
1868 Diplograpsus minimus Carruthers : 74, 130, pi. 5, figs. iaa, b.
?igo6 Climacograptus minimus (Carruthers) Elles & Wood : 191, pi. 27, figs. xa-g.
192 W. CARRUTHERS' TYPE GRAPTOLITES
ORIGINAL DESCRIPTION. " This agrees with D. pristis in general appearance, and
in the form and arrangement of the cells, except that the whole polypary and all its
parts are so very small. Had I met with only a few specimens, I would have con-
sidered it as merely an accidental variety, but I have seen so many, all agreeing in
size, that I cannot doubt that it is a good species, especially as young specimens
of D. pristis early attain their full breadth, and the increase of the polypary is by
addition to its distal end, and not to the size of the already formed hydrothecae,
just as in the living Sertulariadae. About thirty-eight cells to one inch. Loc. Moffat."
FIG. 4. Figures to show Carruthers' clear distinction between his two small species of
diplograptid. a, Copy of original MS drawing of " Diplograptus minimus " which has
appended note "38 to inch ". b, Similar copy of Climacograptus minutus which has notes
" 32 to 40 to an inch, nearly opposite ".
COMMENTS ON USAGE. In his 1868 paper, Carruthers clearly distinguished two
small biserial species and Fig. 4, taken from his notebook, illustrates his ideas of
the differences, one clearly climacograptid, the other what he called diplograptid
(now orthograptid) . His type slab however shows a large number of small, poorly-
preserved rhabdosomes which appear to be almost all climacograptid and Elles &
Wood certainly regarded his species in this light. In his description of the species
(1868 : 130) he called it Diplograpsus minutus although elsewhere, in the explanation
of the plate published in the first part of the paper (1868 : 74), he called it D. mini-
mus, agreeing with his MS notes, and usage of this prior name avoids the homonym
which results from the transference of the species to Climacograptus.
Elles & Wood did not re-figure Carruthers' material, and their specimens, from
the Hartfell Shales (clingani and linearis Zones), reach a breadth of 2 mm.
Carruthers' specimens on the type slab are only about I mm. wide and are associated
with some slender uniserial stipes, which look very like Monograptus sp. (atavus or
acinaces type). It thus seems probable that Elles & Wood's species is not the same
as Carruthers' but it is clear from the foreign references (e.g. Ross & Berry 1963,
Obyt & Sobolevskaya 1964) that it is Elles & Wood's species which is nowadays
W. CARRUTHERS1 TYPE GRAPTOLITES 193
recognized under this name. Carruthers' original notes are of little use since the
only measurements he gives are the thecal numbers per inch. A sketch of " Diplogr.
minimus " clearly shows thecae of an orthograptid type in accordance with his
comparison of it with D. pristis but, as mentioned above, the common form on the
type slab appears to have climacograptid thecae. The species must remain for the
moment in an unsatisfactory state since a proper appraisal of Elles & Wood's form
must await critical re-examination of the Upper Ordovician climacograptids.
TYPE MATERIAL. Q.82, a slab crowded with poorly-preserved specimens.
REVISED DESCRIPTION. The rhabdosome is about i mm. wide and up to 10 mm.
long. Thecal details are obscure.
HORIZON. As noted above, the type slab shows slender uniserial stipes very
similar to monograptids of the vesiculosus and cyphus Zones, in contrast with the
description by Elles & Wood of the species from the Hartfell Shales (dingani and
linearis Zones) .
Climacograptus minutus Carruthers
PL 4, fig. i, ; Figs 3b, 4b
1868 Climacograptus minutus Carruthers : 132, pi. 5, fig. loa.
?i9o6 Climacograptus minutus Carruthers; Elles & Wood : 211, pi. 27, figs. i2a-c.
ORIGINAL DESCRIPTION. ' This is a very minute but well-marked species, never
attaining a greater size then represented on the Plate. There are at the rate of from
thirty-two to forty cells in the space of an inch. Loc. Moffat."
COMMENTS ON USAGE. This species does not seem to have been widely recorded,
probably because of its unsatisfactory nature. Carruthers' original specimens are
of widely differing widths and Elles & Wood based their account of the species on
other material from the British Museum (Natural History). Packham (1962)
revised some of the British Silurian diplograptids but did not discuss this species.
His C. tangshanensis linearis appears to be close to Carruthers' fig. lob (1868).
TYPE MATERIAL. Carruthers' original specimens can be recognized and appear to
be of two different forms. His fig. loa (Q.8o) is here selected as lectotype (PL 4,
fig. i) . The original of fig. lob (Q . 1372) is a narrower form and fits well with
Climacograptus scalaris miserabilis Elles & Wood. Elles & Wood's figured material
(Q . 849) is all somewhat distorted and it is clear that they did not use Carruthers'
original specimens for their description, probably through some confusion with
" Diplogr aptus minutus ".
REVISED DESCRIPTION. Rhabdosome 7 mm. long. 1-2 mm. broad; thecae thirteen
per cm., with large excavations occupying about one quarter of the breadth of the
rhabdosome and about equal to the length of the free ventral wall. The apertures
are nearly opposite each other (as recorded in Carruthers' notes) and not alternate
as stated by Elles & Wood. The proximal end of the type specimen is poorly
preserved but there appears to be a stout virgula which is prolonged beyond the
distal end of the rhabdosome. The type specimen is completely flattened and it is
not possible to make out any details of a median septum.
194 W. CARRUTHERS' TYPE GRAPTOLITES
The uniform breadth of the rhabdosome and the large opposite excavations com-
bine to make this form quite distinct from other British climacograptids. Un-
fortunately the range of variation is not known and since the type specimen has no
associates, its precise horizon is also doubtful. It remains therefore an unsatis-
factory species.
Cryptograptus tricornis (Carruthers)
PL 4, figs. 4-6; Fig. 30
1858 Diplograpsus tricornis Carruthers : 468, fig. 2.
1859 Diplograpsus tricornis Carruthers; Carruthers
1867 Diplograpsus tricornis Carruthers; Carruthers
1868 Diplograpsus tricornis Carruthers; Carruthers
25, fig- 2.
290, pi. i, figs. loa-d.
131, pi. 5, fig. ua, b.
1880 Cryptograptus tricornis (Carruthers) Lapworth : 171, pi. 5, figs. 27a-e.
1908 Cryptograptus tricornis (Carruthers); Elles & Wood : 296, pi. 32, figs. I2a-d.
1908 Cryptograptus tricornis (Carruthers) ; Ruedemann : 443, pi. 28, figs. 1-4.
1913 Cryptograptus tricornis (Carruthers); Hadding : 40, pi. 2, figs. 13-14.
1915 Cryptograptus tricornis (Carruthers); Hadding : 325, pi. 6, fig. 15.
1934 Cryptograptus tricornis (Carruthers); Hsu : 87, pi. 6, figs. I3a-m.
1937 Cryptograptus tricornis (Carruthers) ; Bulman : 5, t-fig. 8.
1945 Cryptograptus tricornis (Carruthers); Bulman : 29, pi. 2, figs. r-8.
1960 Cryptograptus tricornis (Carruthers); Thomas, pi. 6, fig. 69.
ORIGINAL DESCRIPTION. " This species can be readily distinguished by the three
spines which adorn its base, and which are almost always preserved. The central
spine is a continuation of the line of the axis ; it is shorter than the lateral ones . . .
The polypidom is more slender than in D. foliaceus, which in general outline it some-
what resembles. The axis is slender, and produced beyond the other parts of the
fossil. The cell-walls are well marked, extending upwards from the axis to the
boundary of the fossil. Each cell forms a rhomb whose outer border is slightly
indented, giving the boundary of the fossil a faintly serrated aspect. When the
fossil is preserved so as to show the serratures, the spines are so compressed that the
central one is almost or altogether lost. When the spines are well preserved and in
the position described, no traces of the individual cells are discernible ; the boundary
of the fossil is an unbroken line ".
In 1868, Carruthers added " When I described this species I had not detected the
mouths of the cells in those specimens in which they should have been shown on the
upper surface. In more perfectly preserved specimens since obtained these have
been beautifully shown ".
COMMENTS ON USAGE. This species has been widely recognized as it is easily
identified from the original description and figures. Lapworth (1880) discussed the
varying appearance of the thecae at some length and introduced the genus Crypto-
graptus for this and allied species. Hadding (1915) showed by comparison with
Glossograptus that the two stipes were in lateral contact (the monopleural arrange-
ment of Jaanusson 1960) and removed the genus from the Diplograptidae. The
structure of the proximal end was not, however, elucidated until 1938 when Bulman
described isolated specimens from the Balclatchie Limestone and the two different
W. CARRUTHERS' TYPE GRAPTOLITES 195
aspects of the basal spines noted by Carruthers was explained. A number of varieties
have been described, differing mainly in the breadth of the rhabdosome.
DIAGNOSIS. Rhabdosome biserial, monopleural, 2-4 cm. long widest at base
when preserved in lateral view, maximum width 1-8 mm. but typically narrower.
Thecae ten to twelve per cm. Basal spines conspicuous, but short.
TYPE MATERIAL. BM(NH) (^.1299, presented to the museum in 1860, is almost
certainly Carruthers' original slab. It is crowded with specimens up to 35 mm.
long and 1-6 mm. broad but it is not possible to recognize the original figured speci-
mens. Accordingly, one of the better specimens has been selected as lectotype
(PI. 4, fig. 4). The association on the slab includes abundant Corynoides calicularis
Nich. Similar pieces in the Lap worth Collection (Birmingham University) are
labelled " The Cornice, Hartfell " and, although this is not marked on Lapworths'
published map of Hartfell, it would appear from the text that the horizon is lowest
Hartfell, zone of Climacograptus wilsoni.
DESCRIPTION. No detailed description is needed since that in the Monograph
is satisfactory and has been recently supplemented by Bulman's detailed account
of isolated specimens. There seems to be some variation in the thecal count, speci-
mens from Girvan having consistently higher (twelve to sixteen per cm.) counts.
Elles & Wood described the variety schaeferi which Lapworth had figured earlier
and claimed that it was " somewhat wider than the typical form ". Examination
of the material in Lapworth's collection does not confirm this difference but there
seems to be a difference in thecal shape and the basal spines are not conspicuous.
DISTRIBUTION. This species has been recorded from all continents and some of
the more recent records are given in the synonymy. The stratigraphic range was
given by Elles & Wood as Arenig Series (extensus Zone) to Caradoc Series (clingani
Zone), a longer range than any other Ordovician species, and it is possible that
critical examination of a large number of specimens from the lower horizons might
show them to be distinct from the typical form from the early Caradoc. Sherrard
(1954) records it from the zone of P. linearis but no illustration is given to confirm
this.
Monograptus clingani (Carruthers)
PL 5, figs. 1-5
1867 Graptolithus Clingani Carruthers : 369, pi. 2, fig. 8.
1868 Graptolithus Clingani Carruthers; Carruthers : 127, pi. 5, figs, iga, b.
1876 Monograptus Clingani (Carruthers); Lapworth, pi. i, fig. 24.
18760 Monograptus Clingani (Carruthers); Lapworth : 501, pi. 20, figs. sa-c.
non 1897 Monograptus Clingani (Carruthers); Perner, pi. n, figs. 15-17.
1913 Monograptus Clingani (Carruthers); Elles & Wood : 463, pi. 46, figs. na-f.
1951 Monograptus clingani (Carruthers); Bulman : 322, t-fig. 5.
1956 Monograptus clingani (Carruthers); Bondarenko & Keller : 91, t-fig. 2.
ORIGINAL DESCRIPTIONS. 1867. "... a beautiful small species, which at first
I referred to G. millepeda, M'Coy, but that species is certainly the proximal end of
G. Becki, and this differs from it in having a very broad common base, from which
the hydrothecae rise ." 1868. " Polypary, small and arcuate, with a broad common
I96 W. CARRUTHERS' TYPE GRAPTOLITES
canal, and slender somewhat recurved hydrothecae. This beautiful little graptolite
I long supposed to be only the proximal portion of some other species, but the large
number I have met with, all equally perfect, none larger than fig. iga, and many
showing the prolongation of the axis beyond the distal end, together with the great
breadth of the common canal (forming two-thirds of the breadth of the whole
polypary), unlike the early portion or proximal fragment of any graptolite with
which I am acquainted, have induced me to consider it a good species ..."
COMMENTS ON USAGE. Carruthers' descriptions gave no real details of the species.
From the syntypes it appears that two different species may be confused but as
both are young rhabdosomes it is not possible to be sure. The characters of the
proximal thecae are not determinable from the lectotype, chosen by Pfibyl (1948),
which is completely flattened but they appear to be of the priodon type with well
marked hooks to the apertures (PI. 5, fig. 5). The distal thecae are similar. Lapworth
recorded a " Clingani Band ", some 6 inches thick, in his Dobbs Linn sections and
numerous specimens in his collection bear this label. In a recent resurvey of the
section, Toghill (1965) recognized the unit again but recorded M. dingani from a
wider horizon.
TYPE MATERIAL. The specimen (Q.87) figured by Carruthers as fig. iga was
referred to as " type specimen " by Elles & Wood in the explanation of their plate
and the second specimen called " co-type " (Q.8-4), leaving them perhaps of equal
status. Pfibyl (1948) however clearly stated that Carruthers' fig. iga is the lecto-
type which is unfortunate as this specimen has a broader common canal than in
most of the other specimens referred to the species. This may be only a preserva-
tional feature as specimens in the Lapworth collection from DufTkinnel, the type
locality, are all of the common type in which the hooks occupy at least half of the
breadth of the rhabdosome.
REVISED DESCRIPTION. Rhabdosome dorsally curved at the proximal end, dis-
tally becoming more or less straight; widening from an initial breadth of 0-6 mm.
to a maximum of about 1-5 mm. within the first 6 or 7 mm. The thecae are of
priodon type with about one-third of the length involved in the hook which occupies
half of the breadth of the stipe. The shape of the thecae varies a good deal with
the type of preservation but there appears to be no overlap of the thecae (Bulman
1951, fig. 5), the broad prothecal portion occupying the whole of the breadth of the
rhabdosome.
DISTRIBUTION. Lapworth recorded the species in abundance from a 6-inch band
within the sedgwickii Zone at Dobbs Linn, and Marr & Nicholson (1888) similarly
had a M. dingani Band above their convolutus Zone in the Lake District. Elles &
Wood apparently extended the convolutus Zone to include these horizons and also
recorded the species from the gregarius Zone below in their summary table.
Monograptus intermedius (Carruthers)
PI. 5, fig. 6; Fig. 5
1868 Graptolithus intermedius Carruthers : 126, pi. 5, fig. 18.
?i8y6a Monograptus intermedius (Carruthers) Lapworth : 316, pi. 10, figs. loa, d.
W. CARRUTHERS' TYPE GRAPTOLITES 197
non 1913 Monograptus intermedius (Carruthers) ; Elles & Wood : 485, pi. 49, figs. sa-c.
?I952 Spirograptus intermedius (Carruthers); Miinch, pi. 37b, figs, ya, b.
ORIGINAL DESCRIPTION. " Polypary slender; proximal end composed of a slender
canal with distant, isolated, and very small hydrothecae ; adult hydrothecae, short,
triangular, the upper margin of the cell forming an acute angle with the common
canal. About twenty-six cells to an inch. This species differs from G. Nilssoni,
G. tenuis and G. Hisingeri in the form of the cells, and from the last also in the slender
common canal. Perhaps Portlock's figure 6a. pi. 19 of his Report belongs to this
species. Loc. Moffat. "
COMMENTS ON USAGE. It is clear from the specimens in Lap worth's collection
that he included under this name a number of slender forms which do not agree with
Carruthers' type specimen. Elles & Wood used only one slab from the Lap worth
Collection for their redescription of this species and the crowded fragments on it
show thecae which are much more slender than those of the type specimen in which
the thecae are nearer those of M. involutus. Lapworth considered that M. acutus
(Hopkinson 1872) was the same as M. intermedius but Hopkinson quite clearly
differentiated them. Most later workers have, however, apparently relied on Elles
& Wood's account.
TYPE MATERIAL. Holotype, BM(NH) Q.88, Birkhill Shales, Moffat.
REVISED DESCRIPTION. Rhabdosome arcuate, with very slender proximal end,
widening from about 0-3 mm. to a maximum of about i-o mm. in the distal part.
Thecae of the spiralis type, the proximal ones with a slender prothecal portion and
an abruptly widened apertural region, distal ones more uniformly widening, about
ten thecae per cm. The proximal end of the type specimen is poorly preserved but
appears to be comparatively straight so that the rhabdosome is not truly spirally
c
Fig. 5. Sketches of proximal, medial and distal thecae of Monograptus intermedius, holotype,
Q.88, x loapprox.
curved in the way shown by M. communis, etc. On thecal shape, however, Miinch's
assignation of the species to Spirograptus is quite possible. In general shape and
the slender proximal end, the species is close to Spirograptus planus (Barrande) as
figured by Pfibyl (1946, pi. 8, figs. 6-8) but that species rapidly reaches a breadth
GEOL. 17, 4
16
ig8 W. CARRUTHERS' TYPE GRAPTOLITES
of more than 1-5 mm. and the rhabdosome continues to spiral distally. The species
is here retained in Monograptus sensu lato.
DISTRIBUTION. The type specimen is associated with a proximal end of M.
triangulatus-type and seems to be from the zone of M. gregarius. Most of the records
appear to be based on Elles & Wood's description and hence are misidentified so that
no distribution can be given at present.
Diversograptus? capillaris (Carruthers)
Fig. 6a, b.
1867 Rastrites capillaris Carruthers : 368, pi. 2, fig. 10.
1868 Rastrites capillaris Carruthers; Carruthers : 126, pi. 5, fig. 16.
non i8y6a Rastrites capillaris Carruthers; Lapworth : 314, pi. 10, fig. 4.
i8j6a. Monograptus attenuatus Hopkinson; Lapworth : 317, pi. 10, fig. 9.
?i8Q7 Monograptus (Rastrites) gemmatus (Barrande); Perner, t-fig. 26.
1913 Monograptus gemmatus (Barrande); Elles & Wood : 436, pi. 43, figs. 5a-d, ?e.
1952 Monograptus (Psubgenus) capillaris (Carruthers) ; Boudek & Pfibyl : 206, t-fig. 4d-f.
1953 Diversograptus capillaris capillaris (Carruthers) Bou£ek & Pfibyl : 496, 558, pi. i,
figs. 1-3.
ORIGINAL DESCRIPTION. " Common tube very slender, with short isolated tri-
angular hydrothecae, their base of attachment to the common canal as long or longer
than their depth. About sixteen cells to an inch. Loc. Moffat. Richter figures
this specimen in Zeitschr. Deutsch. Geol. Gesellsch.,V, 1853, Tab. xii, fig. 34a, referring
it to R. gemmatus Barr., which is very different, and of which his fig. 34b is a good
representation ".
There has been considerable confusion over this slender species. Carruthers'
original description is not clear and Hopkinson (1872) described M. attenuatus as
another slender species without making detailed comparison with capillaris.
Lapworth took the generic assignment to Rastrites as correct in spite of Carruthers'
statement that the base of the triangular theca was longer than its height. From
his own larger and more detailed collections from Dobbs Linn, Lapworth (i876a)
redescribed M. attenuatus and a " R. capillaris " which is a true rastritid (see Fig. 6d).
Perner (1897) redescribed M . gemmatus (Barrande) which has distinctly hooked
thecae (Fig. 6c) but while he gave only a natural size figure of the holotype he also
gave an enlarged drawing under the same name of a form with simpler thecae.
It was apparently this enlarged figure which Elles & Wood took as typical of M .
gemmatus and accordingly included M . capillaris Carr. and M. attenuatus Hopkinson
in its synonymy. They do not seem to have considered Lapworth's drawings or
specimens of " R. capillaris " (which are clearly distinct from Carruthers' types)
although they refigured his specimens of M. attenuatus. Boucek & Pfibyl (1952)
re-examined the slender monograptids and showed clearly that Carruthers' and Hop-
kinson's species were distinct from M. gemmatus. In the following year they assigned
diversograptid forms to D. capillaris and the species has been widely recorded as
such in Central European work since 1952. Unfortunately British material has so
far provided very few examples of diversograptid rhabdosomes while the German
material (e.g. Manck 1924) is usually poorly preserved as far as thecal characters
W. CARRUTHERS' TYPE GRAPTOLITES
199
are concerned. It is thus doubtful if the equivalence of the British and Continental
specimens can be regarded as proven.
TYPE MATERIAL. Holotype Q . 86, from the Birkhill Shales, Moffat.
REVISED DESCRIPTION. Rhabdosome only known as fragments, up to 5 cm. long,
very slender, maximum breadth 0-4 mm. ; thecae elongate triangular, about seven
per cm., widening gradually from the initial prothecal portion, probably with no
overlap. Apertures of spiralis-type but no spines seen.
DISCUSSION. The type material is very poorly preserved and other slender
British specimens are comparatively rare in collections, generally only found as
small fragments. It is difficult to match these satisfactorily with the German and
Bohemian material which, although more frequently showing complete rhabdosomes,
is usually quite flattened and does not show thecal characters well. The slender
\
FIG. 6. Diversograptus? capillaris Carr. a. Copy from original drawing for Elles & Wood,
pi. 43, fig. 5d. Holotype, Q.86. X5- b. DP capillaris Carr. Two thecae for comparison with
M. gemmatus. Q86. X 10. c. M. gemmatus Barr. Fragment of stipe with 2 thecae showing
hooked apertures. R. maximus Beds, Belcraig Burn, Moffat. BU 304. x 10. d. Rastrites
"capillaris" Lapw. (? = R. spina Richter) Duffkinnell. 611305. Xio.
GEOL. 17, 4 l6§
200 W. CARRUTHERS' TYPE GRAPTOLITES
stipes appear to have been fairly flexible and the shape of the thecae is variable
along the stipe, depending probably on the twisting of the stipe. Although it is
quite clear that D.P capillaris is distinct from M. gemmatus (Barrande), its relation-
ship to M. attenuatus Hopkinson remains doubtful. The specimens which Lapworth
figured as M. attenuatus are obviously close to Carruthers' capillaris but these may
be different from Hopkinson's types which have not yet been traced.
DISTRIBUTION. The type material is associated with Rastrites cf. peregrinus,
indicating a Middle to upper Birkhill age which agrees with the records from the
rest of Europe.
Rastrites maximus Carruthers
PL 5, fig. 7; Fig. ya
i86ya Rastrites maximus Carruthers : 540, fig. 90(6)
1868 Rastrites maximus Carruthers : 126, pi. 5, fig. 14.
i8y6a Rastrites maximus Carruthers; Lapworth, pi. i, fig. i.
1907 Rastrites maximus Carruthers; Tornquist : 15, pi. 2, figs. 28, 29, ?2y, ?3O~33,
?pl. 3, fig. i.
1914 Rastrites maximus Carruthers; Elles & Wood : 494, pi. 50, figs. 6a-d, ?6e.
1941 Rastrites maximus Carruthers ; Pfibyl : 15.
?i955 Rastrites maximus Carruthers; Malinowska : 57, pi. n, fig. 4.
1967 Rastrites maximus Carruthers; Schauer : 184, pi. 6, fig. i.
ORIGINAL DESCRIPTION. " Common tube slender, supporting very large hydro-
thecae at wide intervals. Hydrothecae nearly half an inch long, somewhat enlarged
towards the apex, and furnished at the base with a triangular corneus membrane
extending a short distance up the margin of the cells. About six cells in an inch.
Loc. Moffat. "
COMMENTS ON USAGE. This species has been recorded from a few places in Europe
but appears to be replaced by R. linnaei Barrande in Central Europe. The large
rastritids are usually found in a very fragmentary state owing to the extremely
slender common canal, and isolated thecae cannot always be identified specifically
since the thecal length /interspace ratio appears to be one of the most useful characters
for discrimination. All the large forms, however, appear to occur about the same
horizon and so are useful for stratigraphic purposes.
Although the species has been widely known and figured by Tornquist (1907)
and other authors, no proximal ends appear to have been conclusively demonstrated.
Most of the so-called " young specimens " have short, well-spaced thecae which
may be broken or represent other species such as R. distans. Both Tornquist and
Elles & Wood gave measurements for the first few thecae and the interspaces be-
tween them but examination of a large number of specimens from the Moffat area
and from the Crossfell Inlier, Northern England, shows clearly that the long thecae
of the adult develop within the first centimetre much more rapidly than was generally
thought. Unfortunately few specimens are found with more than a few proximal
thecae and it is difficult to separate R. maximus from R. linnaei and R. distans
prior to the four or five theca stage.
W. CARRUTHERS' TYPE GRAPTOLITES
FIG. 7. Proximal ends of Rastrites spp.
linnaei Barr., BU 1700. Both, x 5 aprox.
a, R. maximus Carr., neotype, BU 1692. b, R.
Table I gives the dimensions of some of these rhabdosomes but the data are as
yet insufficient for a statistical analysis. The positions of theca i and th.2 are the
same in relation to the sicula in the three species, with th2 leaving the common
canal at the apex of the sicula.
R. maximus
Tornquist 1907
Elles & Wood 1913
Neotype, BU 1692
Q.i39oBM(NH)
R. linnaei
BU 1700 (Elles & Wood,
pi. 5 1, fig. ib)
R. carnicus
Seelmeier 1936
TABLE I
length of theca
r
I
2 3
4
I -O
2-7 6-2
1-7
J'5
3'5 5-o
• — •
I-4
5-3 10-2
12-0
i -i
5'3 I0'4
10 +
thecal interspace
. A
I
1/2 2/3 3/4
i -o 1-4 1-7
i • i 3-2 4-8 6-0
1-2 5-0
0-8
0-8
0-6
1-6
4'5
4-0
(0-1—0-3)
TYPE MATERIAL. Carruthers' type specimen cannot be traced in the British
Museum (Natural History) collections and although Q . 1390 bears some resemblance
to Carruthers' figure there are no details of locality or horizon for it. Accordingly
a neotype is here proposed, chosen from the specimens figured by Elles & Wood.
The specimen, 6.11.1692 (Elles & Wood, pi. 50, fig. 6b) is from the Upper Birkhill
Shales of Riskinhope Burn, Yarrow, Peeblesshire. (For stratigraphic details see
LapworthiSyS : 272).
REVISED DESCRIPTION. Adult thecae more than 10 mm. long (maximum observed
1-8 cm.) with interspaces about 5 mm; common canal very slender; base of theca
characteristically enlarged with the proximal side at right angles to the common
canal and the distal side somewhat oblique. Thecal apertures with a well-defined
hook.
DISTRIBUTION. The species is known from many localities in the Moffat area
and was recorded by Elles & Wood from the Lake District and North Wales.
202 W. CARRUTHERS' TYPE GRAPTOLITES
Records from Sweden and Central Europe are doubtful although, as noted above,
it is often impossible to determine specificially the fragmentary large rastritids.
Amongst the Bohemian material in the Lapworth Collection there are no specimens
approaching the typical form in length and spacing of the adult thecae. Recent
records from Belgium (Legrand 1962) and Portugal (Romariz 1962) give no illustra-
tions or dimensions and require confirmation. The specimen figured by Malinowska
(1955) has a broad common canal although the thecae appear to be long enough.
Schauer (1967) has figured a good specimen from Germany.
Cyrtograptus murchisoni Carruthers
PI. 5, fig. 8, 9; Fig. 8
i867a Cyrtograpsus Murchisonii Carruthers : 540, foss. 90, fig. i.
1868 Cyrtograpsus Murchisonii Carruthers; Carruthers : 128, pi. 5, figs. lya, b.
PiSgg Cyrtograptus Murchisoni Carruthers; Perner : 21, t-fig. 28.
1900 Cyrtograptus Murchisoni Carruthers; Elles, pi. 24, fig. 6.
1914 Cyrtograptus Murchisoni Carruthers; Elles & Wood : 505, pi. 51, figs. 3a-c.
?I933 Cyrtograptus murchisoni Carruthers; Boucek : 30, pi. 2, figs. 1-3.
non 1939 Cyrtograptus murchisoni Carruthers; Chang & Sun, pi. i, figs. 4-8.
ORIGINAL DESCRIPTION. " Hydrothecae triangular apiculate, furnished with a
spine. The upper margin of the cell at right angles to the axis, about twenty-eight
cells to the inch. The polypary is considerably incurved at its proximal end, and
as it grows it gradually opens into a larger curve. The branches spring from the
celluliferous surface of the polypary, but as there is no break in the continuity of
the hydrothecae, they must arise from the periderm covering the common canal.
The branches also curve in the same direction as the main portion of the polypary.
Loc. Pencerrig, Builth. I have associated the name of the author of ' Siluria ' with
this remarkable species. The only British specimens I have seen are in the Geological
Museum, Jermyn Street, but among the specimens obtained by the British Museum
from M. Barrande there is a specimen from Listice, labelled G. priodon, which belongs
to this species. "
COMMENTS ON USAGE. Carruthers' figure is somewhat idealized but there is no
difficulty in recognizing the species. The syntypes are not well preserved and do not
show the proximal end well on the large specimens although this may be the result
of damage to the specimens which have at some stage been broken. This in unfor-
tunate as Boucek, in his revision of the Cyrtograptidae (1933), distinguished forms
with excentric proximal coiling from those with central (and more open) coiling.
He had no forms of the first type with secondary branches (a characteristic of mur-
chisoni) but he separated the simply-branched forms (centrifugus and murchisoni
bohemicus) on the proximal ends. The type slabs of murchisoni, however, show
proximal curvature of the centrifugus type and this was also figured by Elles &
Wood.
It is reasonable to separate the simply-branched forms as a subspecies as there
appears to be some stratigraphic value in this (see Rickards 1965) but if the curvature
is also admitted as a diagnostic character, a new name is required for the Bohemian
W. CARRUTHERS' TYPE GRAPTOLITES 203
specimens of " murchisoni ". If the proximal end is regarded as a variable feature
within the species, then there is no good criterion for separating centrifugus from
m. bohemicus and these may both be regarded as junior synonyms of Tullberg's
C. murchisoni var. crassiusculus. Pending a revision with full stratigraphic details
of these Lower Wenlock cyrtograptids, it seems best to restrict the present account
to the redescription of Carruthers' syntypes and the numerous topotypes.
TYPE MATERIAL. Carruthers noted that his types were in the Jermyn Street
Museum and the specimen and counterpart are now numbered GSM 10717-8.
They are from the Wenlock Shale, Pencerraig, Builth, Radnorshire.
REVISED DESCRIPTION. Rhabdosome stout, proximal end forming a helical
spiral with triangular thecae of the spiralis type. First-order cladia curved, arising
at well-spaced intervals (about five to eight thecae between each) and bearing
secondary branches which are generally curved in the same direction. Thecae on
the distal part of the main stipe and on the cladia have a tubular proximal part and
a short, hooked aperture. The change takes place on the main stipe after the third
or fourth cladium. Thecae ten to fourteen per cm.
FIG. 8. Cyrtograptus murchisoni Carr. Poorly preserved proximal end showing excentric
coiling. The most proximal thecae are underneath the later part of the main stipe. GSM
10717. X3.
DISTRIBUTION. As noted above, there is some difficulty in matching Boucek's
description of the Bohemian material with the British types, although Carruthers
recognized the species in material from Listice. The stout, secondarily branched
forms appear near the base of the Wenlock throughout Europe and it appears con-
venient to regard them all as one species. Further geographic variation may be
noticed when well-preserved material from other areas is studied. The species has
been recorded from Canada but so far without confirmatory illustrations. The
form recorded from China appears to be the earlier species, Monograptus spiralis.
Russian records (Obut 1964; Obut, Sobolevskaya & Bondarev 1965) appear to be
of the species group but not the typical form. The relationship of C. murchisoni
to the forms with only first order branches is still unknown but the latter seem
slightly earlier stratigraphically (Rickards 1965) and could well be ancestral. The
absence of cyrtograptids from the succeeding zone of M. riccartonensis suggests that
the later C. rigidus and its allies are not phylogenetically related to C. murchisoni,
since this would involve a reversal of the trend towards more numerous branches.
In my notes on British zones (Strachan 1960) I observed that C. murchisoni was
" apparently unknown outside the type locality ". Since then I have seen many-
branched specimens from the Wenlock of the South of Scotland and Rickards (1965)
has also found numerous specimens in the Lake District.
204 W. CARRUTHERS' TYPE GRAPTOLITES
Dendrograptus lentus Carruthers
i86ya Dendrograptus lentus Carruthers: 541, foss. 90, fig. 5.
1868 Dendrograptus lentus Carruthers: 130, pi. 5, fig. 5.
The specimen described under this name is preserved in the Geological Survey
& Museum, GSM 10691 (and counterpart 10692). The species was noted by Wood
(in Elles & Wood, p. Ivii) as belonging to Clonograptus and a note on the specimens
says " PShineton Shales, Shrops. " Carruthers recorded the species as " Caradoc,
Co. Fermanagh " and the specimens are listed as such in the Survey Catalogue of
1878. However there is a collection number with the drawing of this species in
Carruthers' notes and this appears in the Survey Catalogue of 1864 as " Graptolithus
sp. Cambrian, Wrekin. " There thus is some confusion over this species and since
the name has not been used, so far as I can ascertain, since the original descriptions
it is best regarded as a nomen dubium, and an appropriate recommendation to the
ICZN has been made. If the species were to be regarded as valid, the name lentus
would have priority over the widely used name tenellus, and these fragments are
too poor to be the basis of this well-known species.
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Congress, Pt. 7 : 109-113.
THOMAS, D. E. 1960. The Zonal Distribution of Australian Graptolites. /. Proc. R. Soc. N.S.W.
94 : 1-58, pis. 1-15.
TOGHILL, P. 1965. Stratigraphical variations in the Lower Silurian of the South of Scotland.
Ph.D. thesis, Univ. of Birmingham.
TOMCZYK, H. 1962. [Statigraphical problems of the Ordovician and Silurian in Poland in
the light of recent Studies]. Pr. Inst. Geol., 35 : 134, 4 pis., 13 figs. [In Polish with Engl.
transl.]. Warsaw.
TORNQUIST, S. L. 1907. Observations on the genus Rastrites and some allied species of Mono-
graptus. Acta. Univ. lund., Handl. K. Fysiogr. Sallsk. N.F., 18, 5 : 1-22, pis. 1-3.
TULLBERG, S. A. 1883. Skanes Graptoliter II. Graptolitfaunorna i Cardiolaskiffern och
Cyrtograptusskiffrarne. Sver. geol. Unders. Abh. C, 55 : 1-44, 4 pis. Stockholm.
EXPLANATION OF PLATES
Most of the specimens are in the British Museum (Natural History) and their
numbers bear the prefix Q. Those with prefixes GSM and BU are in the Institute
of Geological Sciences and Birmingham University respectively.
PLATE i
Leptograptus capillaris (Carruthers)
FIG. i. Copy of original figure, Carruthers 1868, fig. ya.
FIG. ?.. Type slab, Q.$o, natural size. Hartfell shales. Hartfell Spa.
Butt. BY. Mus. nat. Hist. (Geol.) 17, 4
PLATE i
GEOL. 17, 4.
PLATE 2
Pleurograptus linearis (Carruthers)
FIG. i. Holotype, Q.848, natural size. Hartfell Shales. Hartfell Spa.
FIG. 2. Copy of page from Carruthers' MS notes showing drawing of type slab and the
original published figures (1858-59) half original size.
FIG. 3. Specimen figured by Nicholson, 1867, fig. i, refigured Elles & Wood, pi. 16, fig. 7.
Q.27, natural size. Hartfell Shales. Hartfell Spa.
Bull. BY. Mus. nat. Hist. (Geol.) 17, 4
PLATE 3
Diceltograptus elegans (Carruthers)
FIG. i. Holotype, Q.850, natural size. Hartfell Shales. Hartfell Spa.
Dicranograptus clingani Carruthers
FIG. 2. Specimen figured Carruthers 1868, fig. 6b, Q.842, natural size. Hartfell Shales.
Hartfell Spa.
FIG. 3. Lectotype, Carruthers 1868, fig. 6a, refigured Elles & Wood, pi. 24, fig. la, Q-55,
natural size. Hartfell Shales. Hartfell Spa.
FIG. 4. Same, x 5.
Dicellograptus moffatensis (Carruthers)
FIG. 5. Page from Carruthers' MS notes showing original drawings and copy of published
figure (1858-59) half original size.
FIG. 6. Type slab, (3.843, natural size. The lectotype, figured by Elles & Wood, pi. 23,
fig. la, is marked with the arrow. Hartfell Shales. Hartfell Spa.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 4
PLATE 3
.
': J -**
:'• £V *'• \ i
\i
s< IF -I:*. ^
PLATE 4
Climacograptus minutus Carruthers
FIG. i. Lectotype, Q.8o, X5. Moffat, horizon and locality uncertain.
FIG. 2. Specimen figured Carruthers 1868, fig. job. (^.1372, as C. minutus but probably
C. scalaris miserabilis E. & W. X5. Moffat, horizon and locality uncertain.
Climacograptus minimus (Carruthers)
FIG. 3. Type slab, Q . 82, natural size, showing poorly preserved biserial graptolites. Horizon
and locality uncertain.
Cryptograptus tricornis (Carruthers)
FIG. 4. Slab with numerous specimens, (^.1299, lectotype marked with arrow, natural size.
Hartfell Shales. Hartfell Spa.
FIG. 5. Drawings from Carruthers' MS notes showing the different appearances of the
rhabdosome half original size.
FIG. 6. Copy of original figures (1858-59) half original size.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 4
PLATE 4
PLATE 5
Monograptus clingani (Carruthers)
FIG. i. Specimen showing distal part of the rhabdosome, BU 1667, figured Lapworth 1876,
fig. 3a, refigured Elles & Wood, pi. 46, fig. nd, natural size. Birkhill Shales. Dobbs Linn.
FIGS. 2 and 4. Specimen figured Carruthers 1868, fig. igb. (3.84, natural size and X5.
MofEat.
FIGS. 3 and 5. Lectotype, figured Carruthers 1868, fig. iga, Q.87, natural size and X5-
Mofiat.
Monograptus intertnedius (Carruthers)
FIG. 6. Holotype, Q.88, figured Carruthers 1868, fig. 18, natural size. Birkhill Shales.
Moffat.
Rastrites tnaximus Carruthers
FIG. 7. Neotype, BU 1692, figured Elles & Wood, pi. 50, fig. 6b, x 2^. Upper Birkhill
Shales. Riskinhope Burn, Yarrow.
Cyrtograptus murchisoni Carruthers
FIGS. 8 and 9. Specimen and counterpart, GSM 10717-8, half natural size. These are the
slabs from which Carruthers made his description and idealized figure. Wenlock Shales. Builth
Bull. Br. Mus. nat. Hist. (Geol.) 17, 4
PLATE 5
«-~ ~ -^ ;
~* * v ^~
, A---
7
PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING
A REVISION OF THE ENGLISH
WEALDEN FLORA, I
CHARALES— GINKGOALES
J. WATSON
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 5
LONDON: 1969
A REVISION OF THE ENGLISH WEALDEN
FLORA, I
CHARALES— GINKGOALES
BY
JOAN WATSON
Pp. 207-254; 6 Plates; 64 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 5
LONDON : 1969
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 17, No. 5 of the Geological
(Palaeontological) series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation
Bull. Br. Mus. nat. Hist. (Geol.).
Trustees of the British Museum (Natural History) 1969
TRUSTEES OF
THE BRITISHfMUSEUM (NATURAL HISTORY)
Issued 21 January, 1969 Price £i 155.
A REVISION OF THE ENGLISH WEALDEN
FLORA, I
CHARALES— GINKGOALES
By JOAN WATSON
CONTENTS
I. INTRODUCTION ......... 210
II. SYSTEMATIC DESCRIPTIONS ....... 214
THALLOPHYTA . . . . . . . . .214
CHLOROPHYCEAE . . . . . . . . 214
CHARALES . . . . . . . . .214
CIRCONITELLA gen. nov 214
Circonitella knowltoni (Seward) n. comb. . . 215
GYROGONA Lamarck 217
Gyrogona medicaginula Lamarck . . . . 217
BRYOPHYTA ......... 218
Musci ......... 218
? Bryophyte leaf ...... 218
HEPATICAE ......... 218
HEPATICITES Walton 219
Hepaticites zeilleri (Seward) n. comb. . . . 219
Hepaticites ruffordi sp. nov. ..... 220
TH A LUTES Walton 220
Thallites valdensis (Seward) n. comb. . . . 221
Thallites catenelloides (Seward) n. comb. . . . 223
PTERIDOPHYTA ......... 224
LYCOPODIALES ........ 224
SELAGINELLA 224
Selaginella dawsoni (Seward) . . . . . 224
EQUISETALES ........ 227
EQUISETITES Sternberg 227
FlLICALES ......... 228
MATONIACEAE ........ 228
MATONIDIUM Schenk ... 228
Matonidium goepperti (Ettingshausen) . . . 228
DlPTERIDACEAE ........ 228
HAUSMANNIA Dunker 228
Hausmannia dichotoma Dunker .... 228
SCHIZAEACEAE ........ 229
RUFFORDIA Seward .... 229
Ruffordia goepperti (Dunker) .... 229
PELLETIERIA Seward ... 232
Pelletieria valdnensis Seward ..... 232
POLYPODIACEAE ....... 235
ASPIDISTES Harris . . 235
Aspidistes sewardi sp. nov. ..... 236
GEOL. 17, 5 J8
2io A REVISION OF THE
UNCLASSIFIED FERNS ....... 237
CLADOPHLEBIS Brongniart . . . . . 237
Cladophlebis longipennis Seward .... 237
Cladophlebis albertsii (Dunker) .... 237
Cladophlebis browniana (Dunker) .... 238
Cladophlebis dunkeri (Schimper) . . . . 238
SPHENOPTERIS Sternberg 238
Sphenopteris ruffordi (Seward) n. comb. . . . 238
Sphenopteris fittoni Seward ..... 238
Sphenopteris fontainei Seward .... 238
LECKENBYA Seward 238
Leckenbya valdensis (Seward) . . . . 238
TEILHARDIA Seward 238
Teilhardia valdensis Seward . . . . . 238
ONYCHIOPSIS Yokoyama 238
Onychiopsis psilotoides (Stokes & Webb) . . 238
WEICHSELIA Stiehler 239
Weichselia reticulata (Stokes & Webb) . . . 240
SPERMATOPHYTA ........ 240
PTERIDOSPERMAE ........ 240
PACHYPTERIS Brongniart 240
Pachypteris lanceolata Brongniart .... 240
GYMNOSPERMAE ........ 242
CYCADALES ......... 242
NILSSONIA Brongniart 242
Nilssonia schaumburgensis (Dunker) . . . 242
BECKLESIA Seward 244
Becklesia anomala Seward ..... 244
Becklesia sulcata sp. nov. ..... 247
GlNKGOALES ........ 248
PSEUDOTORELLIA Florin 248
Pseudotorellia heterophylla sp. nov. . . . 248
III. REFERENCES .......... 251
SYNOPSIS
The English Wealden flora described by Seward (1894, 1895) is here revised using modern
techniques, in particular in the study of the plant cuticles. Much of the work has been carried
out on the existing collections in the British Museum (Natural History), but a number of new
species in the form of isolated leaf cuticles have been obtained by bulk maceration of " coaly-
shales " collected by the author from the Fairlight Clay at Hastings, Sussex.
The present work covers the Thallophyta, Bryophyta, Pteridophyta, Pteridospermae, Cy-
cadales and Ginkgoales. A second part will deal with the Bennettitales and Coniferales together
with a special section devoted to a complete revision of the Wealden Equisetales.
I. INTRODUCTION
English Wealden Flora.
THIS work is a revision of part of the English Wealden Flora, using modern tech-
niques. It has been carried out largely from a study of the Wealden plants in the
British Museum (Natural History). Most of this material was collected from the
coast in the neighbourhood of Hastings, Sussex in the latter part of the last century
ENGLISH WEALDEN FLORA I 211
and belongs to the Beckles, Dawson, Mantell and Rufford Collections. The majority
was collected by Mr. P. Rufford but unfortunately his specimens are merely labelled
" Ecclesbourne, Nr. Hastings " and the exact localities are not recorded. Attempts
were made to find fresh material along the coast between Hastings and Pett Level
but unfortunately no fossils were found. However, material was obtained from
coaly lenses and partings which occur in the Hastings Beds. Bulk maceration of
these " coaly-shales " yielded quantities of spores, isolated leaves and even small
shoots. Two of the species thus obtained are described here, including the first
record of a member of the Ginkgoales from the English Wealden. Similar coaly-
shales from the Wealden in the Isle of Wight yield identical plant fragments.
The flora was previously described by Seward (1894, 1895). His identifications
of the fossils were based solely on macroscopic features and were figured by means of
woodcuts and lithographs. A re-examination of the fossils using modern techniques
shows that many of Seward's determinations are wrong. However, where the type
material of a particular species (usually from the German Wealden) has not been
redescribed or examined it is impossible to say if the identification is truly accurate.
Seward arranged the flora into about seventy-five species. From more than a
dozen of these I was unable to get any significant new information, and unless better
material is found it seems that only a redescription of their gross morphology would
be possible. Most of the ferns come into this category and only a few species are
dealt with here in any detail. There are very few fertile specimens available and
one such fern, Onychiopsis psilotoides (Stokes & Webb), has recently been redes-
cribed by Tattersall (1961). The practice of coating specimens with varnish in the
past has caused the destruction of the cuticle in many specimens and has greatly
hindered revision.
The Thallophyta, Bryophyta, Pteridophyta, Pteridospermae, Cycadales and
Ginkgoales are described in this paper with the exception of the Equisetales. A
recent find of more petrified material belonging to the Equisetales renders necessary
a further revision of the species (see Watson 1964), and this will be published in a
later volume together with the Bennettitales and Coniferales.
Other Wealden Floras
The Wealden floras of Germany, France and Belgium were all described in the
last century in the same way that Seward described the English flora. Since then
little progress has been made and work as detailed as that presented here scarcely
exists. Cuticles have been described for a few species from time to time (Carpentier
1939) but the amount of work to be done remains enormous. The German Wealden
flora in particular should be thoroughly revised as it contains so many type specimens.
A number of English fossils are referred to German species merely on external charac-
ters. When a species under revision can only be compared with the inadequate,
original nineteenth century description of the type an impossible situation arises.
It has hitherto been thought that the English and German floras are more or less
identical but I suspect that this may not be so. In some cases fairly recent work
on foreign material has shown that a number of English specimens previously re-
A REVISION OF THE
f erred to German species are different from the type \Sphenolepis kurriana (Dunker),
Sphenolepis sternbergiana (Dunker), Pseudocycas dunkeriana (Schenk)] and these
were described as new species (Watson 1964). But for the vast majority of species
no such comparison is possible. It is also now known that the German and English
FIG. i. Inferred present extent of Wealden strata. (After Allen 1967)
Wealden are not wholly contemporaneous (see Table I). For these reasons I feel
that it is of little value to produce comparative lists of species for the various locali-
ties. This has been done frequently in the past and perhaps most recently and
comprehensively by Delcourt & Sprumont (1955). Such lists, however, are probably
highly inaccurate and will remain so until there is a wholesale revision of all Lower
Cretaceous fossil floras.
Techniques
Techniques used were of the simplest. With few exceptions the usual method for
bulk macerations was used (see Harris 1926).
Stratigraphical background
The stratigraphy of the Wealden has been described by White (1928), Allen (1954,
1955, 1959, 1960, 1965, 1967), Hughes (1958) and Howitt (1964). Owing to its
alleged relict fauna and flora of Jurassic affinity the English Wealden has on
occasions been assigned to the Jurassic System. It is now, however, generally
ENGLISH WEALDEN FLORA I
213
TETHYAN
BOREAL
South- east
France
North - west
Germany
Southern
England
CRETACEOUS (part)
Barremian
Barremian
Weald
Clay
CRETACEOUS (part)
Hauterivian
Hauterivian
Valanginian
'Upper & Middle
Valdensis
Hastings
Beds
Berriasian
Wealden
Durlston
^ Beds
U
UJ
Upper
Serpulite
Lower Serpulite
~ O£
13 Lulworth
Beds
JURASSIC (part)
Munder
Marls
JURASSIC (part)
Tithonian
Portland
Beds
Eimbeckhausen
Plaltenkalk
TABLE, i — Correlation of N. W. European " Wealden " (from table 4 in Dodson, Rex, Casey &
Allen 1964).
accepted as part of the Cretaceous. Part, or even all, of the Purbeck might also be
Cretaceous. Table I is a correlation table for the Wealden of north-west Europe
recently proposed by Dodson, Rex, Casey & Allen (1964). On the basis of glau-
conite studies the radiometric age of the Jurassic /Cretaceous boundary is estimated
to be about 135 million years. Text-fig, i shows the present extent of the Wealden.
The Fairlight Clays have yielded most of the fossil plants, particularly in the region
of Ecclesbourne Glen. The large number of specimens collected by Mr. P. Rufford
and others has led to the false impression, which has been perpetuated (Gallois 1965),
that plants are abundant in the Fairlight Clays. This is not so: even small frag-
214 A REVISION OF THE
ments are quite rare. It has not been possible to ascertain the exact localities and
horizons at which Rufford collected; nor is it known over what period of time the
collection was assembled. In a brief description of the Hastings Beds by Rufford
(Seward 1894 : xvii) he refers to the Fairlight Clays " yielding, very locally, Ferns,
Cycads and Conifers " and also to " the very limited extent of outcropping fossili-
ferous strata ". In the early years of this century (around 1910) a further important
collection of Wealden plants was made by P. Teilhard de Chardin and Felix Pelletier,
two Jesuit priests who stayed in the area for four years. Their specimens (Seward
1913) were collected mainly in the neighbourhood of Fairlight which is about a mile
to the east of Ecclesbourne Glen. The collection was subsequently divided between
the British Museum (Natural History) and the Hastings Museum. The Hastings
Museum also has a large number of Rufford's specimens, many of which are the
counterparts of the specimens in the British Museum (Natural History). Attempts
at collecting in recent years have yielded nothing to compare with the magnificent
specimens (fronds three feet long) obtained by these earlier collectors.
ACKNOWLEDGEMENTS
This work forms part of a Ph.D. thesis of the University of Reading, supervised
by Professor T. M. Harris, F.R.S. and I thank him for his unfailing patience, kind-
ness and encouragement. I wish also to thank Professor P. Allen for much help and
for permission to reproduce his text-figures.
Some of the work was carried out in the Department of Palaeontology, British
Museum (Natural History), some in the Botany Department, University of St.
Andrews and some in the Herbarium, Manchester Museum where I was given research
facilities for which I am most grateful. Thanks are due to the Keeper of Palaeon-
tology, British Museum (Natural History) for permission to work on the collections.
Tenure of a D.S.I. R. Studentship is gratefully acknowledged.
II. SYSTEMATIC DESCRIPTIONS
Unless otherwise indicated in the text all specimens are in the collections of the
British Museum (Natural History) and are prefixed by V.
THALLOPHYTA
CHLOROPHYCEAE. CHARALES
Organ Genus CIRCONITELLA nov.
DIAGNOSIS. Gyrogonite uncalcified, cutinised; surrounded by five sinistrally
spiralled cells; shape ellipsoidal, circular in transverse section. Apical pole formed
by all five cells; basally ending against a basal plate; without basal cage.
TYPE SPECIES. Chara knowltoni Seward 1894; 13, Text-fig. I.
DISCUSSION. This genus is recognized for the type species which was formerly
included in Nitellites Horn af Rantzien in my Ph.D. thesis of 1964. The diagnosis of
Nitellites, however, requires that size and shape should be very close to that of the
ENGLISH WEALDEN FLORA I 215
Recent genus Nitella but C. knowltoni differs in being somewhat larger and un-
flattened.
Dijkstra (1959) describes an uncalcified gyrogonite which he refers to as " Prae-
chara symmetrica Peck cf. ". I am satisfied, from his diagnosis and photographs
that this fossil is identical with C. knowltoni (Seward). Whilst this species fits into
Praechara Horn af Rantzien (1954) on its general shape and apical structure,
Praechara has hitherto been used only for members of the Characeae, i.e. calcified
gyrogonites which cannot be placed in other well-defined genera.
Circonitella knowltoni (Seward) comb. nov.
PI. i, figs. 1-5 ; Text-figs. 2-6.
1894 Char a knowltoni Seward : 13, text-fig, i.
X959 Praechara symmetrica Peck cf. ; Dijkstra : 15, pi. 2, figs. 20-23.
EMENDED DIAGNOSIS. Gyrogonite uncalcified, ellipsoidal in side view, apical and
basal halves similar, 4007* to 665 JLL long and 380 ju to 550 ju, wide. Spiral cells five,
sinistrally spiralled, about 45 /* wide near equator and showing about twelve con-
volutions laterally. Cells usually slightly concave with distinct intercellular ridges
3 /* to 15 ft wide, sometimes flat and occasionaUy convex. Outer cutinised membrane
(ectosporostine of Horn af Rantzien) marked with coarse but very low and ill-
defined tubercles up to 5 fi diameter. Inner membrane (endosporostine of Horn
af Rantzien) present, delicate and without a patterned surface. Apical pole formed
by ends of spiral cells meeting in a slightly eccentric pattern ; basal pole with rounded
plate 100 /i wide; no cage or basal claws present.
NEOTYPE. ¥.51555.
DESCRIPTION. Numerous black cutinized gyrogonites are embedded in an argil-
laceous matrix, the majority being flattened and presenting a lateral view but a
few being orientated so as to show polar views. A few gyrogonites preserved by
pyritization are completely unsquashed and were obtained by macerating small
pieces of the rock in water. These specimens show that the gyrogonite was originally
completely circular in section and that flattening occurred during preservation.
Whereas most of the flattened specimens have concave spiral cells with intercellular
ridges the solid ones have convex spirals with depressions between and tend to be
somewhat larger. However, one of them (¥.51555) has part of the convex surface
broken away revealing the familiar concave cells with ridges below. These two
surfaces are interpreted as the outer and inner walls of the spiral cells.
Apical poles are not seen nearly so frequently as basal poles but a few undistorted
specimens were obtained. It may be noted that in the material from the Nether-
lands, Dijkstra (1959) also found the apical poles of the gyrogonites less well pre-
served. The tips of the spiral cells tend to be swollen and meet in a slightly irregular
pattern. Many more basal poles are seen with the easily identifiable circular plate
against which the five spiral cells end in a regular manner. Their width near the
pole is the same as at the equator.
Laterally up to twelve convolutions of spiral cells are seen each making more than
two turns. Thus the spiral is so flat that the furrows and ridges appear almost
2l6
A REVISION OF THE
horizontal. As mentioned above the spiral cells are concave in most gyrogonites
but convex in a few specimens. There are also many in which they are flat and
their margins are then nearly, or quite, invisible in the untreated fossil. For
variety in intercellular ridges compare PL i, figs. 2, 4.
The tubercles on the ectosporostine are irregularly arranged about 8 /* apart,
usually about four tubercles across the width of a spiral cell. Many gyrogonites
FIGS. 2-6. Circonitella knowltoni (Seward). 2, 3. Gyrogonites showing apical pole.
Fig. 2, specimen accidentally destroyed. Fig. 3, ¥.51549. x 100. 4. Side view of a
gyrogonite with concave cells, showing the very flat spiral. V. 5 1550. X 75. 5.
Basal pole of a gyrogonite. ¥.3114. x 100. 6. Cutinized wall showing part of three
spiral cells with tubercles on the outer layer. The inner layer of the wall is seen on the
right of the middle cell. Slide V. 5 1551. x 350.
gave no satisfactory cuticles at all and some that did yield cuticles were less clearly
marked. The cuticle is usually reddish-brown in colour but in parts the inner mem-
brane of the wall is seen alone as a straw-coloured homogeneous layer without
markings of any kind. The coronula is not known, nor is the oospore.
Dijkstra suggested that his fossils were originally calcified but became decalcified
during preservation. It is much more likely, however, that the present specimens
were never calcified. There is evidence that both the concave inner wall and the
ENGLISH WEALDEN FLORA I 217
convex outer wall of the gyrogonite cells are composed of organic matter and if my
interpretation of these layers is correct then it seems conclusive. It is clear that
in the specimens with only the inner organic layer present the fine clay matrix is
everywhere pressing against this layer; there is no gap at all such as would be left
if a lime-shell had been dissolved. We would have to suppose that decalcification
occured first, then preservation. This species is preserved in a unique way in speci-
mens from two widely separated localities and it seems unlikely to have been caused
by the same unusual event at two distant places.
The absence of a calcareous shell on the outside of the gyrogonite immediately
suggests Nitellaceous affinities but further study shows that in other features it is
more like the Characeae. I make no assumption about the correct classification of
the plant producing this gyrogonite.
In all the Nitellaceae the gyrogonites are laterally compressed whereas Circonitella
knowltoni is quite round in transverse section as are members of the Characeae. In
studying a number of living Nitellaceae, Horn af Rantzien (1959) found Nitella
hyalina to be the least flattened, a typical specimen being 340 /£ long, 260 fi wide
and 180 ju, thick. He also gives details concerning the average size of Nitella gyrogo-
nites : ' . . . generally speaking the oosporangia (gyrogonites) of Nitella are small, in
most species the length does not exceed 350 fi '. C. knowltoni is much larger,
attaining almost twice this length, although Nitella gyrogonites with larger dimen-
sions do occur, e.g. N. flexilis which reaches 575 fi long (Groves & Bullock- Webster
1917).
Twelve convolutions of spiral cells seen in lateral view is common in the Characeae
but in the genus Nitella the largest number known is eleven, in N. polygyra. A
more usual number in this group is five or six and hence the angle of the spiral is
steeper than in C. knowltoni.
Amongst British Charales the most similar ectosporostine pattern is seen in N.
opaca, although the tubercles are more numerous and larger. Most of the Characeae
have a much smoother granulate surface. The absence of basal claws accords with
Recent species of Nitella.
Apart from Quaternary fossils the only other uncalcified gyrogonites are Nitel-
lites sahnii Horn af Rantzien and Nitellites sp. (Horn af Rantzien 1957) from the
Middle to Upper Jurassic Rajmahal series of India. Both differ from C. knowltoni
in having more in common with the Recent genus Nitella. They are much smaller,
being only 160 /* at the longest, and they are laterally compressed. The spiral
cells show only six to eight convolutions and the ectosporostine has a reticulate
surface.
Genus GYROGONA Lamarck
Gyrogona medicaginula Lamarck
Selected references-
1804 Gyrogona medicagimtla Lamarck : 355.
1829 Chara medicaginula (Lamarck); Lyell : 91, pi. 13, figs. 1-8.
I933 Gyrogona medicaginula Lamarck; Groves : 24, 25, 39.
1965 Gyrogona medicaginula Lamarck; Grambast & Paul : 239, pi. i, figs. 1-4.
218 A REVISION OF THE
REMARKS. Several gyrogonites of this species are present in a piece of pure
siliceous rock (V . 2928) which is included in the Wealden Collections (British Museum,
Natural History). The specimen has the appearance of a large pebble from the
beach and it seems probable that it is not Wealden in age. G. medicaginula is
otherwise only known from the Tertiary (Lyell 1829, Grambast & Paul 1965) and is
found in the Oligocene of the Isle of Wight. The specimen is accompanied by a
note written by Groves and is probably the specimen to which he refers (Groves
T933 : 25) when saying " Fruits collected from a Wealden deposit, Fairlight Glen,
Hastings (Sussex), England . . . are considered by the present writer to be referable
to this species ".
The matrix cannot be matched to anything in the Hastings Beds and the gyro-
gonites of G. medicaginula are much larger than is usual in pre-Tertiary species.
It seems probable, therefore, that the specimen was mistakenly identified as Wealden ;
perhaps washed-up on the beach near Hastings.
BRYOPHYTA
MUSCI
? Bryophyte leaf.
Text-figs. 7-9
Leaf elliptical, nearly 2 mm. long, just over i mm. wide. Margin entire, one end
(possibly both) rounded. No midrib present. Lamina flat, one cell thick, cells
forming about eight longitudinal rows, marginal cells smaller, about as long as broad.
Cell walls thin, not locally thickened at corners; no surface thickenings observed.
MATERIAL. V . 2857, v • 28570, v • 28576.
This small fossil is known from three isolated specimens which are very similar;
one has a counterpart. The above description assumes it is like a Bryophyte leaf
but I have been unable to find a Bryophyte leaf with similarly arranged cells. In
the leafy liverworts the cells tend to be isodiametric or rounded and do not form
longitudinal rows. In Naiadita lanceolata Buckman (Harris 1938) the cell rows
are markedly transverse. In the mosses they are isodiametric to elongated but
they may form longitudinal rows. Transversely elongated cells are very rare in
mosses but they are seen in a few Tortula species. However they have leaves which
are otherwise different.
Ventral scales and rounded appendages of the scales of the Marchantiales have
more or less elongated cells and where rows are recognizable they are transverse,
never longitudinal. In fern ramenta the cells are often in longitudinal rows but are
usually very narrow.
HEPATICAE
The fossils described here are all thalloid and branch dichotomously and some of
them form a rosette-like growth. Some show definite Hepatic features rather than
Algal, but others do not, although they look like Hepatics. Various generic names
have been used for such fossils by earlier authors (e.g. Marchantites Brongniart)
suggesting rather precise affinity. Walton (1925) in giving fine details of such fossils
ENGLISH WEALDEN FLORA I 219
for the first time revised their classification. A fossil with definite Hepatic features
he called Hepaticites. This feature in practice is usually the presence of rhizoids.
Fossils in which no rhizoids were demonstrated he called Thallites. According to
this scheme the genus Marchantites is now restricted to forms showing definite
Marchantialean characters. Thus Marchantites zeilleri Seward was referred to the
genus Thallites Walton (Harris 1942). Further study of this plant has revealed the
presence of rhizoids and it is here included in Hepaticites Walton.
Lundblad (1954) separated from Hepaticites those species showing Marchantialean
ventral scales as Marchantites and those with Marchantialean air-chambers as
Marchantiolites. In the present work the two genera of Walton suffice, but one
species which almost certainly has Marchantialean ventral scales would have been
placed in Marchantites had it been well enough preserved for the facts to be secure.
Genus HEPATICITES Walton 1925 : 565
Hepaticites zeilleri (Seward) comb. nov.
PL 2, figs. 1-3
1894 Marchantites Zeilleri Seward : 18, pi. i, fig. 3.
1942 Thallites zeilleri (Seward) Harris : 397.
1954 Thallites zeilleri (Seward); Lundblad : 408.
EMENDED DIAGNOSIS. Plant thalloid, dichotomously branched, distance between
dichotomies not constant. Thallus about 3 mm. broad, composed of dark, well
defined midrib I mm. wide and delicate lamina probably one cell thick. Margins
entire. Cells of thallus in rows, longitudinal in midrib but inclined at an angle of
about 45° in lamina; cells about 30 fi wide. Near apex midrib tending to divide
into number of divergent dark bands separated by thinner tissue. Numerous
unicellular thin walled, non-tuberculate rhizoids arising on midrib, up to 5 mm.
long and typically 40 p wide.
HOLOTYPE . V . 2330 .
DESCRIPTION. The holotype suggests a rosette shaped thallus in the position of
growth (PL 2, fig. i shows the counterpart). A second specimen shows smaller
pieces of thallus mixed with fragments of Onychiopsis and is more likely to have
been deposited by water.
The thallus branches are delicate towards the tip and the apex is nowhere visible.
Here the midrib is more delicate and seems to divide. In this delicate part longi-
tudinal rows of cells can be seen but elsewhere the midrib is very dark and opaque
and presumably several cells thick. The rows of cells in the lamina are clearly
seen only in a few places and then only the longitudinal walls are preserved.
There is nothing suggesting Marchantialean air chambers or ventral scales. The
rhizoids although clear are only moderately well preserved, unbranched and without
septa; their walls appear to be plain rather than tuberculate. They form groups
arising from the midrib and usually point forwards. A number of detached rhizoid
bunches are present beside the thallus.
COMPARISON. H. zeilleri is the only British Wealden liverwort with a clearly
defined midrib and lamina. Among other fossils it bears a most striking resemblance
220 A REVISION OF THE
to H. laevis Harris (1931) from the Lower Lias of East Greenland. The rows of
cells in that species are more clearly defined and the midrib is 0-5 mm. wider than in
H. zeilleri but otherwise they look the same.
Other similar species are: H. arcuatus (Lindley & Hutton) Harris (1942 : 394,
1961 : 7) which is of similar dimensions and has a well defined midrib, but the lamina
is three cells thick, and there are fewer rhizoids. Thallites yabei (Kryshtofovich
1932, 1938, Oishi 1940, Harris 1942) is a larger plant of which no microscopic details
are known.
CLASSIFICATION. H. zeilleri looks very like Pellia and agrees with it in the cell-
rows of the lamina and in the rhizoids. Plants of the Pellia form are familiar in the
Jungermanniales anacrogynae but scarcely occur in other groups and accordingly
H. zeilleri is placed here.
Hepaticites ruffordi sp. nov.
PL 2, figs. 4, 5
DIAGNOSIS. Thallus branched dichotomously at intervals of about i cm., width
about 3-4 mm. Margins entire, apex probably notched. Greater part of thallus
marked with grooves (considered to be ventral scales) at intervals of 0-5 mm.
Grooves strongly arched, longitudinal in middle region but curving outwards and
then slightly backwards at their ends. Middle region of older part of thallus, about
2 mm. wide, marked by two lines appearing thicker and beyond which grooves do
not extend.
HOLOTYPE . V . 2343 .
DESCRIPTION. The only specimen known is preserved as an impression in fine
sandstone ; a little of the plant substance remaining as a dark stain but this has
cracked and no cells are seen satisfactorily. Faint marks crossing the grooves may
be the impression of cells but there is nothing more. No rhizoids were seen nor any
structure in the lamina. H. ruffordi is tentatively placed in Hepaticites because it is
thought to show a distinctive Hepatic character — ventral scales — even though
rhizoids are not preserved. These ribs, or ventral scales, are of interest since they
are confined to the Marchantiales. Unfortunately the specimen is too poorly pre-
served to be sure that they are indeed scales. I have, therefore, refrained from
including it in Marchantites in Lundblad's sense.
COMPARISON. Marchantites hallei Lundblad (1955) is a liverwort from the Lower
Cretaceous of Patagonia. It is similar in dimensions to H. ruffordi but is known in
much more detail, having two sorts of rhizoids and well preserved ventral scales.
Hepaticites haiburnensis Harris (1961 : 9) is another similar form in which the thallus
extends further beyond the scales.
Genus THALLITES Walton 1925 : 564.
For discussion of Thallites see p. 219. The two species described here are associa-
ted in considerable numbers on certain bedding planes. Seward placed them in
ENGLISH WEALDEN FLORA I 221
Algites thinking them to be marine red algae. Closer study shows that they are
more likely to be terrestrial liverworts though proof is lacking. They are certainly
not marine for the rocks in which they occur are of purely freshwater origin and
indeed the specimens are crowded with the small freshwater lamellibranch, Neo-
miodon. Despite careful study the two forms described here show so few details that
they can hardly be considered to be properly characterized species. However,
they have been described as such and as their names are fairly well known they are
here retained.
Thallites valdensis (Seward) comb. nov.
Text-figs. 12-15
1894 Algites valdensis Seward : 4, pi. i, figs, i, 2.
EMENDED DIAGNOSIS. Plant thalloid; thallus forming rosette of segments branch-
ing dichotomously three or four times. Diameter of whole thallus up to 5-5 cm.;
width of branches typically 4-5 mm., branches ending bluntly. Midrib represented
by median dark line TOO /* wide and dichotomizing 5 mm. behind dichotomy of
thallus. Cells in midrib region not differing from rest of lamina. Dark ribs passing
obliquely across lamina at intervals of about 300 /i. Lamina apparently composed
of one layer of thin walled cells ranging in size from about 15-75 /£.
HOLOTYPE . V . 2857 .
DESCRIPTION. Seven specimens were examined on three blocks of fine-grained
calcareous sandstone. The preservation is poor and splitting of the rock has
resulted also in splitting of the fossil into two parts. The rosette of branches strongly
suggests that the plant has been preserved in position of growth. It accords with a
suggestion made by Walton (1949) for a Carboniferous thallus which he called cf.
Hepaticites. Like Walton's specimen there are several very perfect rosettes and it
looks as though the central and oldest part of the thallus had decayed away before
the plants were buried and preserved. Fragments would be unlikely to drift to-
gether into a neat rosette.
The general appearance of the cells is constant throughout the whole thallus
though they vary considerably in size and shape. They are on the whole thin walled
but additional organic material is present which tends to conceal their walls. The
positions of the median line of the thallus and the ribs are merely marked by a greater
concentration of this dark structureless matter. The central strand is at best faint
and is clear in only two thallus branches. The rest show no central strand at all.
The oblique ribs though not always seen are more widespread. I think that the
pattern of cell-like polygons represents true cells and is not some effect of poor
preservation, for instance the imprint of sand grains on a thin coaly layer. At
best they certainly look like cells and associated with T. valdensis are small frag-
ments of equally delicate plant tissue, also one layer thick but with quite differently
shaped cells. One of these fragments is the small leaf described here as PBryophyte
leaf; another is more like a fern ramentum with very narrow cells. In neither is
there any possibility that the cellular pattern has been imposed by sand grains.
222
A REVISION OF THE
10
kr:*^£^' , -••""'
14
15
FIGS. 7-9. (?) Bryophyte leaf. 7, 8, 9. Leaf showing a single layer of cells. Figs. 8,
9 are part and counterpart. Fig. 7, V. 28576. Figs. 8, 9, ¥".2857. All x 20.
FIGS, 10, ii. Thallites catenelloides (Seward). 10. Part of specimen showing cells.
V.2857. x 100. n. Part of thallus. ¥.2857. x 4.
FIGS. 12-15. Thallites valdensis (Seward). 12. Part of specimen showing cells. ¥.2857.
X 100. 13. Rosette shaped thallus suggesting preservation in situ. V. 28576. x i.
14. A branch of the thallus showing thinner peripheral region and ribs of the thicker
central region. V. 28576 x 4. 15. Apex of branch showing the pattern of diver-
gent ribs. ¥.28576. x 15.
ENGLISH WEALDEN FLORA I 223
No Marchantialean air chambers are visible, nor is there any clear suggestion of
two superimposed layers of cells, though presumably further layers do occur, at
least on the midrib and lateral ribs. No rhizoids are present.
Transfers were made but these yielded no additional information. Chondms
crispus, the red alga with which Seward compared T. valdensis was found to be quite
unlike it in having a much thicker thallus and cells of very different shape.
COMPARISON. T. catenelloides has a thallus of similar texture but is a much smaller
plant. Of the other British Wealden forms Hepaticites zeilleri differs in its clearly
marked midrib and strong rhizoids. H. ruffordi differs in its strong pattern of
arcuate ribs.
Most thalloid liverwort-like plants have a broad and strongly marked midrib.
The following which are of similar dimensions to T. valdensis have only a faint
midrib.
T. umlensis Kryshtofovich & Prynada (1933) from the Rhaeto-Liassic of U.S.S.R.
is rather narrower and has more pointed lobes. No microscopic details are known.
Hepaticites rosenkrantzi Harris (1931, 1937) from the Lower Lias of East Greenland
has no midrib but is two cells thick, the cells tending to be elongated and in rows.
H. amauros Harris (1937), also from the Lower Lias of East Greenland, has a
midrib which is a dark, narrow strip but the thallus is many cells thick.
Thallites catenelloides (Seward) comb. nov.
Text-figs. 10, ii
1894 Algites catenelloides Seward : 7, pi. i, figs, i, 2.
EMENDED DIAGNOSIS. Plant thalloid; thallus dichotomously branched, branches
about 0-5 mm. wide; margins probably entire. Thallus composed of uniform tissue
one cell thick; cells thin walled, diameter about 50/4.
HOLOTYPE. ¥.2857.
DESCRIPTION. Numerous specimens occur on the same blocks as T. valdensis
and in some cases they are closely associated. The specimens of T. catenelloides
vary from small fragments to what are probably whole thalli measuring i cm. across.
They do not form rosettes. The narrow ribbon-like thallus branches frequently
and in parts appears to have an undulating though entire margin. The specimen in
Text-fig, ii is the same as that figured by Seward. His suggestion that the thallus
has constrictions similar to the red alga Catenella seems to be based only on the
appearance of part of this particular thallus. It is not seen in the other specimens
and is probably due to the undulating margin.
The cells of the thallus are identical with those of T. valdensis and the description
for that species will serve for both. Transfers were made but these showed only the
single layer of cells composing the thallus.
COMPARISON. In Catenella opuntia the thallus is several cells thick and has a
surface layer of cells much smaller than those of T. catenelloides, whilst the central
cells are larger and elongated.
GEOL. 17, 5 19
224 A REVISION OF THE
All other Mesozoic hepatics, including the three other English Wealden species,
are much larger than T. catenelloides. The only plants of similar dimensions are
from the Carboniferous. Two rather similar fossils are Thallites willsi Walton (1925,
1928, 1949) and Hepaticites langi Walton (1928). Neither has a definite midrib,
both differ in being more than one cell thick and H. langi has well marked rhizoids.
T. willsi which is more comparable has a rather broader thallus.
PTERIDOPHYTA
LYGOPODIALES
SELAGINELLACEAE
Genus SELAGINELLA Spring
Only one other Selaginella from the Mesozoic, S. hallei Lundblad (19500), is
known in as much detail as the Wealden species described below and referred here
to the living genus.
The genus Selaginellites Zeiller (1906) was originally used for heterosporous fossils
having more than four megaspores in each megasporangium and was thought to
differ in this respect from the Recent genus Selaginella. Subsequently it was used
by Halle (1907) for all heterosporous fossils regardless of the number of megaspores.
Later work proved that living species may produce more than four megaspores in
each megasporangium (Duerden 1929) . Miner (1932) used Selaginellites for isolated
megaspores which would ordinarily have been called Triletes but Harris (1935)
recommended that its use should be restricted to fossil shoots which showed evidence
of being closer to Selaginella than any other genus. Dijkstra (1949) was the first
to place the English Wealden species in Selaginella but this may have been inadver-
tent as he later returned to the use of Selaginellites without comment. I consider
that the Wealden species in no way differs from a typical anisophyllous Selaginella
with the normal number of four megaspores in each megasporangium and it has,
therefore, been referred to the living genus.
Selaginella dawsoni (Seward)
Text-figs. 16-24
1894 Plantae incertae sedis Seward : 20, pi. i, figs. 8, 9.
1913 Selaginellites Dawsoni Seward : 87, text-fig, i.
I9i3« Selaginellites Dawsoni Seward; Seward : 86, pi. 4, figs. 1-7.
1949 Selaginella Dawsoni (Seward) Dijkstra : 19. (name in list).
DIAGNOSIS. Fertile, dichotomously branched shoots; sterile portion of shoot in
part known about i mm. wide, dorsiventral, covered with small overlapping leaves.
Leaves anisophyllous, in four ranks; leaves of upper two rows alternating and
appressed to stem; rhomboid in shape, about 0-5 mm. long; leaves of lower two
rows longer, probably about I mm. long, slightly spreading laterally but pointing
towards apex. Strobilus terminal, unbranched, 2 mm. wide, at least 3 cm. long.
Overlapping sporophylls decussately arranged, apparently all alike and longer than
leaves, about 1-5 mm. Sporangia more or less rhomboid in outline about 0-5 mm.
ENGLISH WEALDEN FLORA I 225
across. Megasporangia fewer but distributed throughout length of strobilus; each
containing four megaspores. Megaspore with rounded dorsal surface, ventral
surface formed by three flattened contact facets ; diameter 285 ^-340 /* (mean 306 /*) ;
height about 15 % less ; tri-radiate ridges very broad, about 30 ju, high, edge rounded,
not divided; length equal to radius of spore; arcuate ridges absent. Ventral surface
crowded with tubercles, circular or oblong and 5-12 fi across ; tubercles also present
on surface of tri-radiate ridges but less prominent. Dorsal surface of spore having
a coarser pattern of widely spaced ribs, irregular in shape, about 4 ju wide and up to
60 [i long; spore surface smooth between ribs. Spore wall easily separable into
two distinct layers, outer 20-30 fi thick, inner about 2 fi thick. Inner membrane
straw-coloured and indistinctly marked with small pits. Substance of wall, in-
cluding tubercles and ribs, light brown in colour and composed of minute granules.
Microsporangia contain numerous microspores which are almost spherical but
with flattened contact facets; average diameter 45 /i (range 38-52 fi}\ tri-radiate
mark fine but distinct, equal to about two thirds of spore radius. Contact facets
seen as three lobes separated by tri-radiate marks not quite reaching notches between
lobes; surface of facets more or less covered with indistinct tubercles, 1-2 ^ in dia-
meter. Spore wall distinctly two layered, together only about 2 ju, thick at facets
but 5-6 fi thick elsewhere. Outside facets both layers uneven in thickness; inner
layer becoming thicker opposite a thin place in the outer. Maximum thickness of
inner layer about 3 /*, outer about 5 fi.
HOLOTYPE . V . 2328 .
DESCRIPTION. The only material available consists of two fragments of shoot
and a few lengths of cone which are present on two blocks of ironstone. Some of
the cone specimens have been much damaged. The two shoots are each about 3 cm.
long and frequently branch dichotomously. For the most part the sterile portion
of the plant is seen merely as a faint outline on the rock, the stem and some of the
leaves being replaced by a white mineral and scarcely any carbonaceous plant material
remains. On only one or two branches can the phyllotaxis be worked out. Here
the mineralized outlines of the dorsal leaves are seen in surface view and the ventral
leaves in side view. This is the portion of shoot seen in Text-fig. 17. Although
the margins of the leaves are shown as entire it is impossible to tell whether they
are correctly represented since the precise margins are unrecognizable. All that is
known of the shape of the ventral leaves is from what they show in side view. They
are seen to have thicker tissue in the centre. The exact arrangement of the dorsal
leaves just behind a dichotomy is uncertain.
The most complete strobilus is 3 cm. long but is probably incomplete, ending as
it does, very sharply at the edge of the rock. Although this strobilus does not have
any attached sterile part, each of the two shoots have bases of similar strobili
attached to one of their branches. Three portions of cones lying close to one another
and parallel indicate that a number of them were borne terminally on the plant.
The sporangial walls are not preserved though the outlines of the microspore
masses are clearly seen. Only one tetrad of megaspores is present in the megas-
porangium.
226
A REVISION OF THE
The sporophylls are seen only at the periphery of the strobilus in side view, their
general shape being unknown.
Slides of detached spores prepared by Dr. W. Chaloner and kindly lent by him
show in most of the megaspores the two very distinctive ornamentations of the
16
17
\
20
24
FIGS. 16-24. Selaginella dawsoni (Seward). 16. Part of a strobilus with microsporangia,
megaspores in outline and sporophylls in side view. V.3I5I. x 10. 17. Part of a
sterile shoot showing anisophylly. Holotype, ¥.2328. x 10. 18. Ventral surface
of microspore showing contact facets, tri-radiate marks and two-layered wall in optical
section. V. 31510. X 500. 19. Dorsal surface of the same spore. V. 31510. x 500.
20. Detail from the same spore showing tubercles on contact facets. V. 31510. x 2500.
21. Ventral face of megaspore. Specimen lost, x 100. 22. Detail of megaspore
showing broad tri-radiate ridge and facet with tubercles. V.3i5iy. x 250. 23.
Single rib from dorsal surface of megaspore. V.3i5ry. x 250. 24. Megaspore in
side view showing dorsal surface, two facets and a tri-radiate ridge. V.3i5ry. X 100.
ENGLISH WEALDEN FLORA I 227
different surfaces and a number of broken specimens show the two separate layers
of the spore wall. The thin inner layer is easily removed and mounted in glycerine
jelly without further treatment. Very brief maceration of the the thick outer
coat renders it transparent before mounting. Some of the megaspores do not show
the two distinctive sculptures quite so clearly as the figured specimens. In these
the dorsal ribs are not so coarse and well defined. Lack of abundant material
prevents more detailed study of this variation.
Some of the mounted microspores remain in tetrads but most of them occur singly.
The layers of the wall are clearly seen in optical section, their irregular adjacent
surfaces closely interlocking, the one being at its thickest where the other is thinnest.
The pits in the outer wall while most numerous on the dorsal side also occur on the
ventral side but not on the facets.
By squashing the spores the thin walled facets may be detached and their surface
more easily studied. In some specimens the tuberculate pattern covers the whole
surface of the facet but is only marginal in others.
COMPARISON. Only one other Mesozoic Selaginella shoot with cone has been
described, namely 5. hallei Lundblad (1950^) from the Rhaetic of Sweden. The
sterile shoot is similar in size but with slightly larger and more spreading leaves.
Like S. dawsoni it is anisophyllous, but the leaves are known in more detail. The
strobilus of 5. hallei is much shorter and has sporophylls which are described as
spirally arranged but may equally well be described as in alternating whorls of
about four. There are four megaspores in each megasporangium. Selaginellites
polaris Lundblad (1948) is a Selaginelloid cone from the Trias of Greenland. It is
much stouter than 5. dawsoni (7 mm. wide) and has numerous megaspores in each
megasporangium.
Selaginella dichotoma Velenovsky (1931) is a sterile shoot from the Cretaceous of
Bohemia with more spreading leaves.
Selaginellites nosikovii Kryshtofovich & Prynada (1932) is a sterile shoot from the
Jurassic of Siberia which also has larger, more spreading leaves than 5. dawsoni.
Among living species 5. sanguinolenta is very similar in size and leaf arrangement,
but the spores are different the megaspores being of similar size but with much more
prominent tubercles on the dorsal surface.
5. emmeliana possesses strikingly similar megaspores to those of 5. dawsoni with
the same shape and sculpturing both on the dorsal and ventral surfaces. S. emmel-
iana, however, possesses other megaspores which show a great deal of variation
both in size and surface ornament. Not all have different dorsal and ventral pat-
terns, some having the coarse ribbed pattern on both surfaces whilst others have the
fine tuberculate pattern on both. The foliage of 5. emmeliana is quite different from
that of 5. dawsoni.
EQUISETALES
Genus EQUISETITES Sternberg
REMARKS. The external morphology of E. lyelli has been fully described by
Allen (1941). Additional information about this species (Watson 1964) was obtained
228 A REVISION OF THE
from some petrified fragments found by Mr. P. Whybrow (British Museum, Natural
History). Very recently Mr. Whybrow has found much more extensive petrified
material and in addition new compression material is available with easily prepared
cuticle. It will, therefore, be necessary to make a complete revision of the species
using all available material, including that described by Allen (1941).
Descriptions of all three species of Wealden Equisetites (Seward 1894) will be
included in a later volume.
FILICALES
Family MATONIACEAE
Genus MATONID1UM Schenk 1871 : 219
Matonidium goepperti (Ettingshausen)
1852 A lethopteris goepperti Ettingshausen : 8, pi. 7, fig. 7.
1894 Matonidium Gopperti (Ett.); Seward : 63, text-figs. 7, 8.
1913 Matonidium goepperti (Ett.); Seward : 89, text-fig. zC, pi. 14, fig. 3a.
1961 Matonidium goepperti (Ett.) ; Harris : 112, text-figs. 37, 38. (see also for earlier references).
Matonidium goepperti (Ett.) has been fully revised by Harris (1961) using abundant,
well preserved material from the Yorkshire Jurassic. The English Wealden material
is scanty and ill-preserved but agrees in all available characters with the Yorkshire
material and I regard the two as specifically identical.
Family DIPTERIDACEAE
Genus HAUSMANNIA Dunker 1846 : 12
Hausmannia dichotoma Dunker
PI. 3, figs, i, 2
1846 Hausmannia dichotoma Dunker : 12, pi. 5, fig. i ; pi. 6, fig. 12.
1894 Dictyophyllum Roemeri Schenk; Seward : 140, text-figs. 16, 17.
1906 Hausmannia dichotoma Dunker; Richter : 18, pis. 3-6.
1913 Hausmannia pelletieri Seward : 89, pi. 14, figs. 1-3.
1961 Hausmannia dichotoma Dunker; Harris : 126, text-fig. 43.
This species is widespread in the Lower Cretaceous of Europe and less common in
the Jurassic. It has been revised by Harris (1961) who gives an emended diagnosis
based on Richter's material (Lower Cretaceous ; Germany) and on Yorkshire Jurassic
specimens. The leaves of H. dichotoma, both sterile and fertile, are quite well known
but nothing is yet known of the sorus. The English Wealden material consists of a
few imperfect sterile leaves which add nothing to our knowledge of the species.
The best examples are figured in PI. 3, figs, i, 2.
ENGLISH WEALDEN FLORA I 229
Family SCHIZAEACEAE
Genus RUFFORDIA Seward 1894 : 75
Ruffordia goepperti (Bunker)
PL 4, figs. 1-7; PL 5, figs. 1-4; Text-figs. 25-31.
Selected references :
1843 Cheilanthites Gopperti T>\mk.&c : 6.
1894 Ruffordia Gopperti (Dunker) Seward : 76, pi. 3, figs. 5, 6; pi. 4; pi. 5, figs. 1-5.
1894 Ruffordia Gopperti (Dunker) var. latifolia Seward : 85, pi. 6, figs, i, ia.
1910 Ruffordia Goepperti (Dunker) ; Seward : 350, text-fig. 260 A-C. (figures repeated from
Seward 1894).
1913 Ruffordia goepperti (Dunker); Seward : 91, text-fig. 2A. (drawing of a spore).
1921 Ruffordia Goepperti (Dunker); Halle : 10, pi. 2, figs. 1-8.
1958 Ruffordia goepperti (Dunker); Couper : 109, pi. 17, figs. 4-6. (photographs of spores).
1966 Ruffordia goepperti (Dunker); Hughes & Moody-Stuart : 274, pi. 43, figs. 1-12. (spores).
EMENDED DIAGNOSIS. Frond triangular, tripinnate to quadripinnate ; rachis
slender. Pinnae triangular or ovate; primary pinnae nearly opposite; secondary
pinnae increasingly alternate. Sterile and fertile parts distinct. Sterile pinnules
variable in width, decurrent; narrow pinnules linear, apices mucronate or acute,
venation Coenopterid; wider pinnules ovate, Sphenopterid, with small marginal
tooth at each vein ending. Fertile pinnules reduced in length but not breadth;
more widely outspread, not decurrent, margins undulating, apices bluntly rounded.
Sporangia borne singly on underside of pinnules without indusium ; typically twelve
on a pinnule, arranged in two longitudinal rows. Annulus transverse, apical,com-
posed of about fifteen to twenty-five long narrow cells radiating from a central plate ;
extending at least to middle of sporangium. (Remaining parts of sporangium
unknown, spore number uncertain). Spores rounded-triangular with convex sides,
typically 45 /£ equatorial diameter (range 35 fi to 60 //.), tri-radiate cracks almost
reaching corners. Apical parts of facets smooth, rest of facets and whole of distal
surface covered with ribs separated by grooves. Ribs forming three sets, each set
starting on contact face and extending on to distal surface ; in each set ridges nearly
parallel. Ridges of a set joining near tri-radiate cracks and also joining, though
less regularly, along continuation of tri-radiate cracks onto back of spore. Smooth
apical parts of facets as thick as ribs.
DESCRIPTION AND DISCUSSION. Ruffordia goepperti (Dunker) is represented by a
large number of sterile specimens which show a wide range in the form of the pin-
nules, the extremes of which are shown in PL 4, figs. I, 4. The specimens with
broadest pinnules were designated R. goepperti var. latifolia (Seward 1894) but all
are now included in R. goepperti (Dunker) as there seems to be intergradation,
though the possibility of the widest forms belonging to a different species is not
excluded. There are three fertile specimens amongst the English material (V.2i6o
V . 2295, V . 2192) . Two are completely fertile (V . 2160 and V . 2192) whilst the other
(¥.2295) has basal fertile pinnae and a sterile top (PL 5, fig. i). Specimens ¥.2192
and ¥.2295 are preserved in the characteristic red ironstone and only the spore
masses, and sometimes the sporangium annulus, are well preserved, the rest of the
230
A REVISION OF THE
31
FIGS. 25-31. Ruffordia goepperti (Dunker). 25. Broad form of pinnules showing vena-
tion. V.2357. x 8. 26. Fertile pinnules showing undulating margins. Broken
lines indicate positions of sporangia. V.2i6o. x 15. 27. Sporangium showing thick-
ened walls of annulus. V.22Q5. x 100. 28. Spore in equatorial view showing
widely spaced ribs and shapes of ribs in section, i.e. rounded, flattened, grooved. Slide
from V.2i6o, x 750. 29, 30. Proximal and distal surfaces of the same spore showing
wide ribs closely spaced. Slide from V. 21 60, X 750. 31. Spore with narrow ribs and
wide grooves. Slide from V. 2 1926, x 750.
ENGLISH WEALDEN FLORA I 231
frond being only a faint impression on the rock. This is a typical condition of the
plants in this matrix but it is sometimes useful for details of venation (PL 4, fig. 7).
Specimen V . 2160 in a soft, fine, grey siltstone matrix is much better preserved though
the frond is heavily varnished and badly cracked. Text-figure 26 shows the rounded
lines of the fertile pinnules with a lobed appearance. The specimen presents the
upper surface of the frond and the sporangia below are seen only in faint outline.
A balsam transfer revealed no details of the sporangia.
The spores of R. goepperti (Dunker) were inadequately figured by Seward (1913,
text-fig. 2A) using ¥.2192 and by Couper (1958, pi. 17, figs. 4-6) using Seward's
slide preparation. Recently the spores have been studied in detail by Hughes &
Moody-Stuart (1966) using ¥.2192 and Seward's slide mentioned above. I have
found that V . 2192 yields spores which are somewhat different from those of V . 2295
and V.2i6o. The spores agree in size range and general form but differ in details
of sculpturing. The spores from V . 2295 and V . 2160 have ribs (muri) from 2 /* to 5 /*
wide separated by furrows (lumina) 0-5 [i to 2 fi wide but the ribs of any particular
spore are always wider than the furrows and the majority of spores have ribs about
3 ju, wide and furrows I /£ wide (Text-figs. 28-30). The ribs vary in profile and may
be rounded, flattened or indented; this variation is seen in Text-fig. 28. Spores
from ¥.2192 have the sculpture in almost exactly the reverse proportions (Text-fig.
31), the ribs being i ju, to 2 /i wide and spaced 2 fi to 5 /£ apart. Hughes & Moody-
Stuart (1966) give a detailed diagnosis of these spores and numerous figures. I
have prepared spores from various parts of ¥.2192 but I have been unable to find
any with broad ribs. Neither ¥.2295 nor V.2i6o yielded spores with narrow ribs
and I could find no suggestion of gradation between the two types of spores.
The identity of ¥.2192 is not in doubt, however, because in all other respects it
agrees perfectly with the other two specimens. It is closely similar to the basal
pinnae of ¥.2295 and also has undulating pinnules as in V.2i6o.
Fertile specimens of R. goepperti (Dunker) from Manchuria were figured by Halle
(1921) including photographs of the sporangia, each represented by a well preserved
annulus. This annulus is certainly an apical ring of the Schizaeaceous type. Halle's
photographs show the long annulus cells radiating from a small circular space or
" plate ". This suggests that the annulus cells in Text-fig. 27 do not show their
full length, although he does report cases where the " central area seems to be much
wider than others ". Halle removed what he considered to be a whole sporangium
and gives three countings in the region of 400. I have removed what appeared to be
whole sporangia from the balsam transfers and obtained various counts under 100.
I regard my counts as highly suspect as it was difficult to be sure that the spore
masses were whole sporangia. One cannot know how reliable Halle's count is but
he reports that " in some cases it was possible to remove the whole contents of a
sporangium more or less intact ". In living members of the Schizaeaceae the spore
output is 128 or 256. Hughes & Moody-Stuart (1966) regard the spores of Halle's
material as of a different type from those found in R. goepperti (Dunker) from the
English Wealden. However, his photographs of sterile and fertile fronds appear to
be very similar to the English specimens. One of them (pi. 2, fig. 2) shows short,
undulating, fertile pinnules. Halle's photograph of the spores (pi. 2, fig. 8) is a
232 A REVISION OF THE
poor one and considering the spore differences in the English specimens of R.
goepperti (Dunker) I have retained the Manchurian material within the species
until it can be proved otherwise. Certain details of the sporangium in the diagnosis
above are based on Halle's account.
Among living genera of the Schizaeaceae, R. goepperti (Dunker) most resembles
Anemia with its distinct and reduced fertile pinnae. The sterile frond of Anemia
adiantifolia is very similar in form to the broader specimens of R. goepperti whilst
the narrow pinnules closely resemble Anemia cuneata (Bower 1926, text-fig. 440).
The sculptured spores of the genus Mohria are of exactly the same type as R.
goepperti (Dunker). This type of spore also occurs in the other Wealden Schizaea-
ceous fern, Pelletieria valdensis Seward and a comparison is included in the descri-
tion of that species.
Genus PELLETIERIA Seward 1913 : 91
Pelletieria valdensis Seward
PI. 6, figs, i, 2; Text-figs. 32-40
1913 Pelletieria valdensis Seward : 91, text-figs. 2B, 3, 4; pi. 12, figs. I2a, I2&; pi. 14, fig. 5.
1958 Pelletieria valdensis Seward ; Couper : 109.
1966 Pelletieria valdensis Seward; Hughes & Moody-Stuart, pi. 44, figs. 1-8; pi. 45, figs. 1-7.
EMENDED DIAGNOSIS. Known only from fertile specimens. Frond at least
quadripinnate ; consisting of slender axis branched repeatedly, with no lamina except
at tips of ultimate branches. Tips bearing groups of fertile pinnules; each pinnule
tightly rolled, completely enclosing the sporangia. Each rolled pinnule more or
less spherical, 2 mm. across, enclosing at least ten sporangia; sporangium elongated,
sessile or shortly stalked, wall composed of small elongated cells, longitudinally
orientated. (Annulus unknown). Spores rounded-triangular with tri-radiate cracks
of medium length ; surface sculptured with a pattern of flat ribs 3-7 /* wide separated
by narrow grooves 0-5-2 ju, wide.
LECTOTYPE. ¥.2329. Selected Hughes & Moody-Stuart 1966.
DESCRIPTION. P. valdensis Seward is represented by several specimens all of
which are fertile. Fragments of pinnae and detached fertile " capsules " are
abundant but there are only two assembled groups of pinnae, specimens ¥.2368
and ¥.2329. Both of these syntypes were figured by Seward (1913, text-figs. 36, B';
pi. 12, figs. I2«, b.) and ¥.2329 (Text-fig. 32) has recently been selected as Lectotype
by Hughes & Moody-Stuart (1966). After I picked away the matrix ¥.2368 is
now a better specimen, more extensive and better preserved than ¥.2329 (PI. 6,
figs, i, 2, Text-fig. 39).
The exact form of the fertile " capsules " is far from fully understood. A number
of isolated " capsules " were dissolved out of the rock in HF but revealed nothing
more about their structure than is shown in the Lectotype which clearly shows the
folded nature of the pinnules (Text-fig. 32). A number of specimens show hairs in
the matrix where " capsules " have been removed (Text-fig. 32) but such hairs could
ENGLISH WEALDEN FLORA I
233
FIGS. 32-36. Pelletieria valdensis Seward. 32. Lectotype showing folded " capsules ",
hairs and cell directions (indicated by dotted lines). V.232Q. x 6. 33. Small
specimen with three ribs on the " capsules ". V. 51846. x 6. 34. Largest assem-
bled shoot showing dissected form of frond. ¥.2368. x 3. 35. Spore showing
thickness of wall (indicated by dotted line) V. 51841. x 750. 36. Eroded "cap-
sule " with one sporangium showing cells of the wall. Others showing sporer V. 51847.
X 35-
234
A REVISION OF THE
not be found on any of the specimens dissolved out. Certain specimens show the
axis continued as a rib over the back of the rolled fertile pinnules and this rib is
branched in ¥.51846 (Text-fig. 33). The pinnules of ¥.51846 appear less tightly
rolled than others and might well prove worth sacrificing to a balsam transfer.
That this specimen is fertile is not in doubt as spores can be seen through a broken
part at the top.
Because the sporangia are completely enclosed they could only be studied in
certain eroded " capsules " which show sporangia in various attitudes (Text-fig. 40) .
38
FIGS. 37—40. Pelletieria valdensis Seward. 37, 38. Proximal and distal surfaces of the
same spore. Slide from V. 2329, x 750. 39. Equatorial view of spore showing widely
spaced ribs. Slide from V. 51841, x 750. 40. Eroded " capsule " with folded
pinnules in section and spores in sporangia. ¥.51840. x 35.
ENGLISH WEALDEN FLORA I 235
Attempts to isolate a sporangium failed because the plant material crumbles very
easily into small fragments: thus the spore number is not known. However, on
such a sporangium as in the centre of Text-figure 40 around fifty spores can be counted
on the surface but because only one layer can be seen it seems probable that the
contents are at least twice this figure. One sporangium seen in surface view (Text-
fig. 36) shows the cells of the wall, small, elongated and longitudinally orientated;
although there is no indication of the annulus.
The spores of P. valdensis Seward have recently been redescribed by Hughes &
Moody-Stuart (1966) using Seward's figured slide preparation from ¥.2329 (1913)
and new preparations from ¥.2329. They found the size range to be 52-98 p with
80 % of specimens between 63 jn and 84 ju, maximum diameter. My preparations
from V . 2329 and V . 21841 agree well with these figures but V . 2368 yielded somewhat
larger spores, 56-115 /* (100 measured) with 75 % between 75 ju and 105 fi. Hughes
& Moody-Stuart (1966) have diagnosed, described and figured the spores in detail
and there is nothing more to add.
COMPARISON. The ribbed spores of P. valdensis Seward are typically Schizaea-
ceous and very similar to those of various living and fossil species. However, the
form of the fertile fronds cannot be matched ; the living members of the Schizaeaceae
merely having curled margins protecting the sporangia. The only comparable
structure I know is in the species Onoclea sensibilis Linn, where the fertile pinnules
are narrow and so tightly inrolled as to completely enclose the sporangia.
The spores of P. valdensis are similar in form to those of Ruffordia goepperti,
particularly the form with the broad ribs and narrow grooves, but they are quite
easily distinguishable on size difference alone. The upper limit found for the dia-
meter of R. goepperti was 60 /* and the lower limit for P. valdensis 63 /*. A most
distinct difference is the thickness of the exine; in R. goepperti about 4 /* (including
the thickness of the ribs) whilst in P. valdensis (Text-fig. 35) it varies from about
5 /£ to 15 fji. This variation depends upon the spaces between ribs being open or
closed and is dealt with in some detail by Hughes & Moody-Stuart (1966 : 280-282) .
A further difference is that the ribbed pattern in P. valdensis extends on to the
contact facets whilst in R. goepperti the facets are smooth.
Family POLYPODIACEAE
Genus ASPIDISTES Harris, 1964 : 181
The type species of this genus, Aspidistes thomasi Harris, is the only other Mesozoic
fern I know with the indusium superum and other aspidioid characters. Although
various diagnostic characters of Aspidistes are not known for Aspidistes sewardi
sp. nov. the form of its indusium and spores give it a good measure of agreement.
The slight difference in spore number (A. thomasi has forty-eight )is not important.
However, further details of the sorus and sporangium may well show it to be generi-
cally distinct. Various other genera exist whose names suggest close affinity with
Aspidium or Polystichum but are based only on sterile specimens.
236 A REVISION OF THE
Aspidistes sewardi sp. nov.
PL 6, figs. 3, 4; Text-figs. 41-43
1894 Weichseliamantelli (Brongniart) Seward : 121.
DIAGNOSIS. (Based on single fertile pinna V.2i73«). Pinnules arising from
upper surface of rachis at an angle of 50-60°; pinnules 2 mm. long, 1-5 mm. broad,
apex rounded; joined to each other at base by a narrow web; margin of pinnules and
web rolled. Groups of three sori borne on under surface of each pinnule ; two larger
adjacent to the rachis. Each sorus with a thick peltate indusium covering about
twelve sporangia arranged in a circle. Sporangium containing thirty-two spores
(sporangial wall unknown). Spores trilete, smooth walled, thickened at corners;
narrow bands of thickening flanking triradiate crack. Arms of triradiate crack
nearly reaching corners, sometimes bifid at ends ; diameter 60-75 /£.
HOLOTYPE. V . 21730:, the only specimen.
DESCRIPTION. The single specimen (¥.2173^) is 6-5 cm. long and was formerly
referred to Weichselia mantelli (= W. reticulata Stokes & Webb) by Seward (1894)
its form being very similar to the smaller pinnules of that species. However its
abaxial surface is uppermost and on moistening with paraffin the sori are clearly
visible to the naked eye (PI. 6, fig. 4). It is certainly not Weichselia reticulata for
fertile material of that species is included in the Bommer Collection and it is quite
different (correspondence with Dr. K. L. Alvin).
Unfortunately the specimen is heavily impregnated with glue and the substance
of the fossil is badly crumbling, making it very difficult to examine the details of
the sorus. The water soluble gum swells rather violently when wet and lifts the
fossil fragments off the surface of the rock. Careful picking with a fine needle re-
vealed a certain amount of information. The circular indusium appears to be quite
thick and is probably attached by a central stalk; details of its cellular structure
could not be seen. The sporangia are distinct pear-shaped masses, about twelve
in number, in a ring under the indusium and sometimes a few under the rolled margin
of the pinnule. They were easily removed intact to count the spores. Of the twelve
sporangia macerated and counted nine contained thirty-two spores and three others
contained twenty-nine, thirty, and thirty-one. The sporangial wall was not seen
either in the untreated sporangium or during maceration.
Among the Wealden ferns there are few species of sterile fronds to which this
specimen could belong but it is similar to one described as Dichopteris delicatula
Seward (1913 pi. n, figs. 6a, b). Possibly further specimens, sterile or fertile, are
among the Weichselia reticulata material but nothing is known of its venation and
it is not always clear in Weichselia. Furthermore any specimen with the adaxial
side upwards would easily be missed.
COMPARISON. Amongst living genera A. sewardi is most similar to Polystichum
agreeing in the form of the indusium and the spore output. However all living
species of the Aspidieae have wedge-shaped spores whereas both A. sewardi and
A . thomasi Harris have trilete spores.
ENGLISH WEALDEN FLORA I
237
42
44
43
FIGS. 41-43. Aspidistes sewardi sp. nov. 41. Three pinnules showing shape, rolled mar-
gins and sori. V.2i73a, x 15. 42, 43. Two spores. Triradiate crack in Fig. 43 with
bifid ends. Fig. 42, slide ii from V. 2173(3, x 750. Fig. 43, slide i from V. 21730, x 500.
FIG. 44. Nilssonia schaumburgensis Dunker. Base of leaf . Specimen L. 5532 Manchester
Museum, x 5.
A . sewardi is named after Sir Albert Charles Seward who first described the English
Wealden flora.
UNCLASSIFIED FERNS
The following are ferns, described by Seward (1894), to which I can add little or
nothing.
Cladophlebis longipennis Seward
1894 Cladophlebis longipennis Seward : 89, pi. 9, figs, i, la.
Cladophlebis albertsii (Dunker)
1894 Cladophlebis Albertsii (Dunker) Seward : 91, text-fig. 9; pi. 8.
238 A REVISION OF THE
Cladophlebis browniana (Dunker)
1894 Cladophlebis Browniana (Dunker) Seward : 99, pi. 7, fig. 4.
1913 Cladophlebis Browniana (Dunker) Seward : 95, pi. 8, figs, i, 2.
Cladophlebis dunkeri (Schimper)
1894 Cladophlebis Dunkeri (Schimper) Seward : 100, pi. 7, fig. 3.
1950 Cladophlebis (Klukia) dunkeri (Schimper); Radforth & Woods : 780, pi. i.
The various English Wealden specimens, referred by Seward to the four species
above, appear to be indistinguishable from each other and Cladophlebis longipennis
Seward is particularly ill-defined. Indeed Seward himself later wrote (1913) of the
" difficulty — or indeed, impossibility — of separating Cladophlebis browniana and
Cladophlebis dunkeri ". Modern treatment of Canadian material referred to C,
dunkeri (Schimper) by Radforth & Wood (1950) shows the fern to have Schizaeaceous
sporangia and spores, though the latter are scarcely distinguishable in the photo-
graph.
Sphenopteris ruffordi (Seward) comb. nov.
1894 Acrostichopteris Ruffordi Seward : 61, pi. 6, fig. 3.
Sphenopteris fittoni Seward
1894 Sphenopteris Fittoni Seward : 107, pi. 6, fig. 2; pi. 7, fig. i.
Sphenopteris fontainei Seward
1894 Sphenopteris Fontainei Seward : 106, pi. 7, fig. 2.
Leckenbya valdensis (Seward)
1894 Nathorstia valdensis Seward : 145, pi. 7, fig. 5; pi. 9, figs. 2, 2«.
1894 Leckenbya valdensis (Seward) Seward : 384.
1895 Leckenbya valdensis (Seward); Seward : 225.
1911 Gleichenites cycadina (Schenk); Seward : 664.
Teilhardia valdensis Seward
1913 Teilhardia valdensis Seward : 96, pi. n, figs, ja-gb.
Genus ONYCHIOPSIS Yokoyama 1889 : 26
Onychiopsis psilotoides (Stokes & Webb)
1824 Hymenopteris psilotoides Stokes & Webb : 423, pi. 46, fig. 7.
1827 Hymenopteris psilotoides Stokes & Webb; Mantell : 55, pi. i, figs. 3a, 3b; pi. 3, figs. 6, 7;
pi. 3*, fig. 2.
1828^ Sphenopteris Mantelli Brongniart : 50.
1828 Sphenopteris Mantelli Brongniart; Brongniart : 170, pi. 45, figs. 3-7.
1833 Sphenopteris Mantelli Brongniart ; Mantell : 241.
1894 Onychiopsis Mantelli (Brongniart) Seward : 41, pi. 2, fig. i; pi. 3, figs. 1-4.
1894 Onychiopsis elongata (Geyler) ; Seward : 55.
1905 Onychiopsis psilotoides (Stokes & Webb) Ward : 155, 506, 518, pi. 39, figs. 3-6; pi. in,
fig. 4; pi. 113, fig. i.
ENGLISH WEALDEN FLORA I 239
1910 Onychiopsis Mantelli (Brongniart) ; Seward : 378, fig. 280, A, B.
1913 Onychiopsis mantelli (Brongniart); Seward : 96.
1961 Onychiopsis psilotoides (Stokes & Webb); Tattersall : 349, pi. 12, figs. 1-3.
DISCUSSION. This species was of world wide distribution in the Lower Cretaceous
and its remains are known from England, Germany, France, Belgium, Portugal,
Russia, Canada, U.S.A., Egypt, South Africa, India and Japan. Although the
species has been described numerous times, including fertile parts (Tattersall 1961),
it is still imperfectly known. Spores were readily obtained but resemble those of a
number of families and do not help in classification. Details of sporangia are still
unknown ; though some fertile pinnules have a lumpy surface suggestive of sporangia.
However the whole surface is covered by an indusium-like sheet of elongated cells
and maceration leaves no trace of sporangial walls.
I have been unable to add usefully to our knowledge of this species and consider
that the figures of Seward (1894) and Tattersall (1961) cannot be improved upon.
In the light of a recent review of this species by Sukh Dev (1965) with an exhaustive
synonomy I have given only those references which include the important name
changes or refer to the English material.
Genus WEICHSELIA Stiehler
Weichselia reticulata (Stokes & Webb)
PI. 6, fig. 5
Selected references :
1824 Pecopteris reticulata Stokes & Webb : 423, pi. 46, fig. 5; pi. 47, fig. 3.
1827 Pecopteris reticulata Stokes & Webb; Mantell : 56, pi. 3, fig. 5; pi. 3*, fig. 3.
18280 Lonchopter is Mantelli Brongniart; 60.
1828 Lonchopteris Mantelli Brongniart : 369, pi. 131, figs. 9, 10.
1894 Weichselia Mantelli (Brongniart) Seward : 114, text-figs. 12, 13.
1899 Weichselia reticulata (Stokes & Webb) Ward : 651, pi. 160, figs. 2-4.
1919 Weichselia reticulata (Stokes & Webb); Florin : 305, figs, i, 2-5.
1965^ Weichselia reticulata (Stokes & Webb); Reyman6wna : 16, text-fig, i; pi. i, figs. 7-11.
1965 Weichselia reticulata (Stokes & Webb) ; Sukh Dev : 73, figs. 7, 8.
DISCUSSION. This widespread Lower Cretaceous species has recently been re-
vised by Reymanowna (19650) and Sukh Dev (1965). Most of the English specimens
are mere fragments of pinnae, many of them fusainized, but there is one large,
assembled frond in the collection. This is figured by Seward (1894, pi. 10, fig. 3.)
and shows the characteristic habit of the plant very well.
The lower epidermis of W. reticulata (Stokes & Webb) was first figured by Florin
(1919) using Belgian material and Sukh Dev (1965) has figured the epidermis from
English Wealden specimens. Reymanowna (1965) gives figures of stomata together
with photographs of indusia, but the fertile material of W. reticulata (Stokes &
Webb) has yet to be fully described. None of the English material is fertile.
Sukh Dev (1965) gives extensive references to older work so that only selected
references have been cited above.
GEOL. 17, 5 2O
240 A REVISION OP THE
SPERMATOPHYTA
PTERIDOSPERMAE
Genus PACHYPTERIS Brongniart
1828 Pachypteris Brongniart : 49 (Name only) .
1829 Pachypteris Brongniart : 166.
1964 Pachypteris Brongniart ; Harris : 121 (Emended diagnosis).
Pachypteris lanceolata Brongniart
PL i, fig. 6; Text-figs. 45-47
1828 Pachypteris lanceolata Brongniart : 50 (Name only).
1829 Pachypteris lanceolata Brongniart : 167, pi. 45, fig. i.
1895 Dichopteris sp. cf. D. laevigata (Phillips) Seward : 184, pi. 12, fig. 6.
1964 Pachypteris lanceolata Brongniart; Harris : 137, text-figs. 55-58.
MATERIAL. V. 3245
DESCRIPTION. The following description is based on the only English Wealden
specimen known. It is a single pinna, 8 cm. long, i cm. wide at the widest part,
with two rows of pinnules, leathery in texture. The rachis, up to 700 /* wide, is
very prominent on the underside of the pinna. The pinnules arise alternately,
usually slightly overlapping but not completely divided and are up to 6 mm. long
and 3 mm. wide, narrowing slightly but attached to the rachis by a broad and
decurrent base. The pinnules are typically oval and blunt ended. Pinnules at the
apex are smaller and less divided; the rachis is not so prominent here. The venation
is visible on the lower side of some pinnules projecting slightly above the surface,
but concealed in most, the veins consisting of a midrib with a few laterals given off
at a fairly low angle. The midrib is broad and situated above the middle of the
pinnule where it joins the rachis. The margins of the pinnules are not thickened.
The upper epidermis lacks stomata and has a thick cuticle showing isodiametric
cells with smooth walls [and without pits. These cells are irregularly arranged
except at the margin where they are elongated and tend to be in longitudinal rows,
the cells here having thicker anticlinal walls. The lower cuticle is thinner, the cells
of the lower epidermis being thin walled, isodiametric and irregularly arranged,
except at the margin and in the midrib region. The position of the midrib is indi-
cated by a region of elongated, thicker walled cells in longitudinal rows. The margin
is like that of the upper epidermis. Stomata are numerous on the lower side but
absent from the midrib and marginal regions. They are slightly sunken, irregu-
larly orientated and distributed and are almost always separated by ordinary
epidermal cells. The guard cells are thinly cutinized except along the edges of the
stomatal aperture where there is a thicker rim; the aperture is elongated. Subsidiary
cells, usually six in number, form a ring and are generally smaller than the other
epidermal cells. They have thicker cuticle on the periclinal walls which forms a
ring round the stoma but are not thus thickened adjacent to the stomatal pit.
Encircling cells are occasional and unspecialized. Trichomes are absent.
DISCUSSION. The preservation is good though the surface is somewhat cracked
and because of the shortage of material available only a little could be macerated.
ENGLISH WEALDEN FLORA I
241
However, it was established that the specimen is attached to the rock by its upper
surface and that whilst the lower cuticle is complete most of the upper is missing,
only a narrow strip remaining around the edge of the pinnules. Thus very little
upper cuticle has been examined and it is not known whether it had any stomata
nearer the middle of the pinnule. Only a few of the pinnules show the positions of
the midrib and the laterals on the underside, and at best they are indistinct. An
attempt was made to see the venation by transmitted light during maceration but
this failed. The prepared cuticle, whilst showing the position of the midrib by the
elongated cells and lack of stomata, gives no indication of the position of the lateral
veins.
47
FIGS. 45-47. Pachypteris lanceolata Brongniart. 45. Part of pinna showing venation.
¥.3145. x 4. 46. Reconstructed transverse section of pinna showing prominent
rachis on lower side. X4- 47. A single stoma. The darker stippling indicates the ring of
thicker cuticle. Slide V. 31456. x 700.
The surface of the specimen viewed in reflected light shows very clearly the cells
of the lower epidermis with a divergent trend in relation to veins. In the frag-
mentary cuticle preparations this trend is not apparent and the cells appear hap-
hazard in arrangement. One very prominent cell was seen, the nature of which is
obscure though it is certainly not stomatal. It is possibly a trichome base though
no similar cells could be identified with certainty.
COMPARISON. A careful comparison was made with specimens of Pachypteris
lanceolata Brongn. from the Yorkshire Jurassic (Harris 1964). The Yorkshire
material is known in much more detail, numerous specimens being available some
of which are complete leaves. A notable feature of P. lanceolata is its wide range
in form with intergradation between the extreme types. These include forms with
small oval pinnules, others with small narrow pinnules and much larger forms with
242 A REVISION OF THE
lanceolate pinnules. Forms just like the present specimen occur. The cuticles of the
various forms although basically the same show a certain variety in details such as
number and size of stomata. Some have a large number of stomata close together
while in others they are more widely spaced. The cuticle of the present fossil shows
more widely spaced stomata than most of the specimens attributed by Harris to
P. lanceolata but some of the latter approach the Wealden form very closely. These
include specimens from Boulby Alum Quarry, Shale 2B ; Whorlton, Scugdale, Rank
Crag Soft Jet and Eston Moor, Beacon Quarry. It should be pointed out that these
are not ordinary specimens but merely cuticle fragments obtained by maceration
and identified on their microscopic characters.
As in the Wealden fossil, the stomata of the Jurassic P. lanceolata avoid the midrib
and margins and are indistinguishable in structure. There is also very close simil-
larity in size and shape of the ordinary epidermal cells, indeed some of the cuticle
fragments of the two fossils could scarcely be told apart.
What little is known of the venation of the Wealden form simply affords more
evidence of the similarity between these two fossils and consequently I have placed
the Wealden specimen in the same species.
GYMNOSPERMAE
GYGADALES
Genus NILS S ONI A Brongniart
Nilssonia schaumburgensis (Dunker)
Text-figs. 44, 48-52
Selected references :
1846 Pterophyllum schaumburgense Dunker : 15, pi. i, fig. 7; pi. 2, fig. i; pi. 6, figs. 5-10.
1871 Anomozamites schaumburgense (Dunker) Schenk : 231, pi. 33, figs. 1-9.
1890 Nilssonia cf. schaumburgensis (Dunker) Nathorst : 5, 9, 13, pi. i, figs. 6-ga.
1895 Nilssonia schaumburgensis (Dunker) ; Seward : 53, text-fig. 3.
1905 Nilssonia schaumburgensis (Dunker); Fontaine in Ward : 303, pi. 72, figs. 17-21.
EMENDED DIAGNOSIS. Leaf linear; width usually 3-5 mm., rarely up to 12 mm.;
length unknown but at least 9 cm. Apex rounded and notched; base tapering.
Midrib 1-2 mm. wide, depressed on upper surface, depression not narrowing towards
apex. Lamina entire or more or less divided. Surface of lamina flat or slightly
convex upwards. Veins fairly prominent, projecting above and below; arising from
midrib at an angle between 70° and 80°; concentration of veins typically three to
four per mm.
Cuticle of moderate thickness but anticlinal walls often obscure. Marginal cells
of leaf and pinnae elongated parallel to edges and having thicker and more distinct
anticlinal walls. Cells in main part of lamina tending to be elongated almost at
right angles to edge, parallel with veins. Upper cuticle without stomata or tri-
chomes, positions of veins scarcely distinguishable. Lower cuticle not showing
positions of veins clearly but cells with slight tendency to be more elongated over
veins. Stomata scattered between veins; small, inconspicuous and scarcely sunken.
ENGLISH WEALDEN FLORA I
50
243
52
FIGS. 48-52. Nilssonia schaumburgensis (Dunker). 48. Leaf, slightly divided, showing
form of apex. Position of rachis indicated by dotted line or solid line where lamina is
broken and rachis is exposed. V.3762. x 2. 49. More divided leaf showing the
veins. Position of rachis shown by dotted lines. V.2I7I. x 5. 50. Cuticle showing
elongated cells of leaf edge in centre, lower surface to the left, upper surface to the right.
Slide from ¥.1436. x 300. 51. Lower cuticle showing a resin body, a stoma and
two single-celled trichome bases. Slide from V.2I7I**. x 300. 52. Single stoma
showing exposed guard cells. Slide from ¥.3762. X 700.
244 A REVISION OF THE
Guard cells exposed; 30-50/4 long and about 15 {i wide, surrounded by several
unspecialized and ill-defined subsidiary cells. Unicellular trichome bases scattered
over whole lower surface; thickly cutinized except for central circle. Resin bodies
occasionally seen, about 50 fi across. Reproductive organs unknown.
DESCRIPTION. This small Nilssonia is represented by more than a dozen speci-
mens from the English Wealden. They show a good deal of variety in the form of
the lamina, some being entire, others slightly and irregularly divided, others re-
gularly and completely divided almost to the midrib. Only two specimens are
more than 6 mm. wide, V .2ijia which is 12 mm. wide (Seward 1895 : 53, text-fig.
30) and V.373I which Seward labelled as Taeniopteris beyrichii var superba. Both
agree with the others in their cuticle. All but one specimen have the adaxial surface
exposed and in these the midrib is seen only when the substance of the lamina is
broken away (Text-fig. 48) but in all specimens its position is indicated by a depres-
sion of the lamina. This depression retains its width to the apex of the leaf. V . 3731
is the only specimen with the lower surface uppermost and is the largest specimen,
9 cm. long (incomplete) and u mm. wide, tapering to 4 mm. near the base. The
rachis is 2-5 mm. wide whilst in most specimens it is little more than I mm.
Resin bodies are rare; they are not seen on the surface of the fossils and only three
were seen in macerated cuticle fragments.
The stomata are particularly difficult to distinguish and were identified with
certainty on only a very few pieces of lower cuticle. The stoma in Text-fig. 52
is the best example seen and shows a slight surface thickening in the middle region
of the guard cells.
DISCUSSION. Nilssonia schaumburgensis has been identified by numerous authors
from several countries. In all cases there is very little detail on which to compare the
various accounts. From the few details available, i.e. size, shape and vein concentra-
tion the English material agrees well with the type-material described by Dunker
(1846) and the specimens figured by Schenk (1871), Nathorst (1890) and Fontaine
in Ward (1905) .
COMPARISON. N. revoluta Harris (1964) from the Jurassic (Upper Deltaic) of
Yorkshire is the only other species I know which is as small as N. schaumburgensis.
The leaves are similar in size and are entire; also the vein concentration is similar,
up to thirty per cm. Another similarity is that the veins are scarcely shown in the
upper cuticle, however they are strongly indicated in the lower cuticle. Rather differ-
erent characters shown by N. revoluta include conspicuous cell outlines with nodular
thickenings and bulging subsidiary cells.
Genus BECKLESIA Seward 1895 : 179
Becklesia anomala Seward
Text-figs. 53-56
1895 Becklesia anomala Seward : 179, pi. 14, figs. 2, 3.
EMENDED DIAGNOSIS. Leaf simply pinnate, probably much longer than broad
(whole leaf unknown), consisting of a slender rachis bearing two lateral rows of
ENGLISH WEALDEN FLORA I
245
long straight pinnae. Rachis 6 mm. wide, pinnae arising at intervals of approxi-
mately i cm., at an angle of about 40°. Pinnae at least up to 10 cm. long, 3 mm.
wide, narrowing to about 2 mm. near rachis (apex unknown), rounded in transverse
section; margins of pinnae straight. Lower side of pinna having broad, flat midrib
53
54
55
FIGS. 53-56. Becklesia anomala Seward. 53. A single stoma. One hypodermal cell
is seen on the right indicated by a single line; Slide from V. 2361 a. x 700. 54. A
piece of cuticle from a non-stomatal region of the leaf, showing a four-celled trichome
base. Slide from V. 23610,. x 200. 55. An unsquashed portion of the pinna showing
the two narrow stomatal grooves on the lower side. V.236ia. x 10. 56. A recon-
structed section of a pinna, x 10.
246 A REVISION OF THE
flanked by two very narrow, shallow stomatal grooves with lateral non-stomatal
regions about as broad as the midrib; upper surface flat but some pinnae showing
numerous raised trichome bases.
Cuticle about 2 IJL thick in non-stomatal regions, very delicate in stomatal grooves.
Ordinary epidermal cells of upper and lower surface large, thin walled and arranged
in longitudinal rows ; very varied in shape, either square, rectangular or quite irreg-
ular; typically 50 fi broad. Cells over midrib and at edges of pinna tending towards
diamond shape. Anticlinal walls cutinized, usually straight but sometimes slightly
sinuous. Presence of a hypodermis occasionally indicated, particularly near trichome
bases. Trichome bases mostly occurring on upper surface but sometimes also in
non-stomatal regions of lower surface ; roughly circular in surface view, about 60-70 fi
diameter, composed of up to four small, thick-walled cells tending to overlap other
epidermal cells. Trichome bases not present on all pinnae. Stomata longitudinally
orientated and scattered in grooves about 45 fi wide. Guard cells thinly cutinized,
about 45 /* long with six to seven subsidiary cells surrounding and slightly overlapp-
ing them; guard cells and subsidiary cells slightly sunken in shallow stomatal pit,
surrounded by thicker walled cells, inner walls of which form thickly cutinized pit
rim. Ordinary epidermal cells between stomata much smaller than those in non-
stomatal regions.
HOLOTYPE. V. 23610.
DESCRIPTION. This species is described from two specimens both of which are
incomplete leaves with no complete pinnae. Whilst most of the pinnae have been
squashed during preservation, two fragments are so preserved that they retain their
original shape and thickness. One is about i mm. thick and shows particularly
well that the stomatal grooves are shallow, that the midrib is flat, and that the margin
is rounded in section.
Much of the plant substance is missing from the two specimens and what remains
is very cracked, so that all cuticle preparations are small pieces. Cuticle from the
non-stomatal regions was easily obtained but it was only with considerable difficulty
that preparations of the stomatal grooves were made. All grooves are filled with
matrix and have extremely delicate cuticle which disintegrates even when most
carefully macerated. The stoma shown in Text-fig. 53 is from the best preparation
obtained in which, unfortunately, all the stomatal pits contain debris which obscures
some of the details.
DISCUSSION. While the characters of this species are distinctly Cycadalean it
is not closely comparable to any other genus, living or fossil. It is thus convenient
to retain the original generic name. The stomatal grooves are much narrower
than in any other Cycad. In Cycas stomata are present over the whole of the lower
surface except the midrib and margins. The trichome bases of B. anomala are
similar to those in Cycas revoluta (Pant & Mehra 1962, text-fig. 330) but in that
species they are two-celled.
Amongst fossil Cycads Paracycas cteis Harris (1964) from the Jurassic (Lower
Deltaic) of Yorkshire is similar in size but differs in such features as wider stomatal
bands and irregularly arranged stomata.
ENGLISH WEALDEN FLORA I
Becklesia sulcata sp. nov.
247
PI. 2, fig. 6; Text-figs. 57-58
DIAGNOSIS. Pinnae up to 2 mm. wide (length unknown), with two narrow longi-
tudinal grooves on lower side; stomata confined to these grooves. Grooves so
placed that the central non-stomatal region is twice as wide as lateral non-stomatal
regions. Margins smooth and entire. Cuticle moderately thick but much thinner
in stomatal grooves. Non-stomatal region between grooves composed of cells in
longitudinal rows, transverse anticlinal walls usually oblique, sometimes slightly
FIGS. 57, 58. Becklesia sulcata sp. nov. 57. Cuticle from a stomatal groove showing
stomata and papillae. Holotype, V. 51524. x 300. 58. Cuticle of upper side.
V. 51524. x 300.
sinuous. Cells of other non-stomatal regions not arranged in rows, very varied in
shape and irregular in arrangement; walls often slightly sinuous. Guard cells
extremely thinly cutinized, except for slight thickening around stomatal aperture.
Guard cells about 50 /* long, not at all sunken, irregularly orientated but majority
more or less longitudinal. Subsidiary cells very variable, not distinguished from
other ordinary epidermal cells in groove, all of which are irregular in shape and size,
smaller than epidermal cells of non-stomatal regions. Subsidiary cells often shared
by adjacent stomata. Some of these cells bearing large, hollow, globular or elongated
papillae; usually one such papilla near each stoma.
HOLOTYPE. ¥.51524.
DESCRIPTION. This species is known only from cuticle fragments which are
assumed to be pieces of pinnae although there is no direct evidence that they are
248 A REVISION OF THE
from a pinnate leaf. However, it is so similar to Becklesia anomala Seward that I
have placed it in the same genus. The main similarities are in such features as
width of pinnae, position and width of stomatal grooves and form of ordinary epider-
mal cells.
COMPARISON. Becklesia sulcata lacks the multicellular trichome bases of B.
anomala, has thinner more exposed guard cells and has large papillae.
If the form of ordinary epidermal cells is any criterion in considering the affinities
of these little known plants, these irregular yet characteristic cells are almost exactly
the same as in many living Cycads e.g. Cycas revoluta.
GINKGOALES
Genus PSEUDOTORELLIA Florin 1933 : 142
EMENDED DIAGNOSIS. Leaves entire, linear to elliptic; apex rounded, narrowing
at base. Two or more longitudinal veins formed by dichotomies chiefly in basal
part, ending blindly. Stomata confined to lower side, always longitudinally orien-
tated; haplocheilic, monocyclic or incompletely amphicyclic; guard cells sunken,
strongly cutinized. Subsidiary cells four or more. Epidermal cells with straight
or faintly sinuous outlines.
The new fossil described here is at least the ninth species of this genus to be des-
cribed and it now seems necessary to emend the generic diagnosis as the original
diagnosis is too narrowly defined for present purposes, based as it was on the single
species P. nordenskioldi Florin. The simplified, broader diagnosis above will better
serve to admit such a fossil as the new English Wealden species with its wide range
of leaf form and scattered stomata.
TYPE SPECIES. Pseudotorellia nordenskioldi Florin 1933.
Pseudotorellia heterophylla sp. nov.
PI. 6, figs. 6, 7; Text-figs. 59-64.
DIAGNOSIS. Leaves entire, showing wide range of size and shape from narrow and
needle-like to broad and elliptic. Dimensions of holotype 12 mm. long and 3 mm.
wide ; longest leaf known 3 cm. long and i mm. wide ; other leaves ranging between
these two. Apex unknown. Petiole unknown.
Veins varying in number from two in narrowest leaf to eight in broadest, formed by
dichotomies near base of leaf. Veins may converge near apex but end blindly.
Interstitial ducts sometimes present between veins.
Cuticle thick and leathery. Stomata present on one side of leaf only, scattered
over whole surface (presumably lower), not avoiding veins; longitudinally orientated,
typically 80 [i long. Ordinary epidermal cells elongated, arranged in longitudinal
rows, sometimes with elongated, thickened ridge along surface. Surface of cuticle
frequently sculptured.
Guard cells sunken beneath subsidiary cells, strongly thickened at common
junction wall with overlying subsidiary cells. Subsidiary cells six, usually papillose,
ENGLISH WEALDEN FLORA I 249
papillae projecting over stomatal aperture. Stomata usually not upsetting se-
quence of cell rows, subsidiary cells continuous in their lines of ordinary epidermal
cells. Sometimes small subsidiary cells coinciding with outer edge of guard cells,
forming ring or partial ring of more specialized subsidiary cells. Encircling cells
rare. Guard cells sometimes having long polar appendages.
Upper epidermis without stomata, composed of longitudinal rows of elongated
cells with fairly thick walls, sometimes in " packets ", often having finely sculp-
tured cuticle. Positions of veins not indicated on upper surface. Edges of leaf
smooth with fairly thick rim of cuticle. Cuticle of all forms of leaf closely agreeing;
only minor differences in size of stomata, degree of thickening and form of polar
appendages of guard cells.
HOLOTYPE. ¥.51525.
DESCRIPTION. A considerable number of whole and broken leaves were obtained
by bulk maceration of the same coaly material which yielded Becklesia sulcata.
Besides these a few of the larger needles were picked off specimen V.2222 in the
British Museum (Natural History) collections. Several fragments were also iden-
tified in washings from the core of a borehole drilled near Winchester. Most of
the linear leaves were found only in short lengths whilst the large elliptical leaves
were almost whole. However, on none of these is an apex or petiole present.
The petiole presumably had one vein which dichotomized at the base of the leaf to
give the two main veins which may further give off branches on their inner sides in
the lower part of the leaf. The veins are then parallel along the length of the leaf
but may converge slightly near the apex (Text-figs. 61, 62.) although they all appear
to end blindly. The interstitial ducts which are often present between veins are
of unknown nature but are possibly resin ducts or even subsidiary veins.
A number of leaves have what appear to be abortive stomata. One of these is
seen in Text-fig. 63. No guard cells are developed but a ring of papillae surround
the " guard cell mother-cell ". A few other " freak " stomata were seen where
the guard cells were misplaced under the stomatal pit. Only one transversely
orientated stoma was seen.
The differences in polar appendages of guard cells may be compared in Text-figs.
63, 64. Text-fig. 64 shows a stoma with the longest and most complex appendages
projecting well beyond the ends of the guard cells. In Text-fig. 63 the appendages
are quite short and pointed or scarcely developed at all. Also shown in Text-fig. 63
are some stomata with the subsidiary cells following the line of the guard cells as
mentioned in the diagnosis.
COMPARISON. P. heterophylla shows a much wider range of leaf form than other
species. P. nordenskioldi (Nathorst) described by Florin (1933), P. minuta (Nathorst)
by Lundblad (1957), P. ensiformis (Heer) and P. crassifolia (Prynada) by Doludenko
(Vachrameev & Doludenko 1961) all have the greatest width of their leaf situated
towards the apex. P. grojecensis Reymanowna (1963), P. ephela (Harris 1935) and
the elliptic leaves of P. heterophylla have the greatest width of the leaf situated
in the middle region. P. angustifolia Doludenko and P. longifolia Doludenko
250
A REVISION OF THE
(Vachrameev & Doludenko 1961) both have long linear leaves of similar dimen-
sions to some forms of P. heterophylla.
A common feature of the Pseudotorellia cuticle is the presence of median ridges of
cuticle on the epidermal cells. P. heterophylla, P. nordenskioldi, P. minuta, P.
ephela and P. ensiformis all have these ridges. In all, the epidermal cells are similar
in form, longitudinally elongated. The packets of cells seen in the upper epidermis
59
61 r?
60 ;
FIGS. 59-64. Pseudotorellia heterophylla sp. nov. 59-62. Various forms of leaf showing
veins. Figs. 60-62 showing interstitial ducts. Fig. 59, ¥.51526, Fig. 60, V. 51527,
Fig. 61, Holotype, V. 51525, Fig. 62, V. 51528. All x 10. 63. Cuticle showing stomata,
abortive stoma and ridges on the epidermal cells. V. 51529. x 150. 64. Stoma with
long polar appendages Slide from V. 2222. x 300.
of P. heterophylla (PI. 6, fig. 6) are surpassed in development by those seen in P.
grojecensis. In that species the packets consist of up to five cells and form a con-
spicuous pattern at right angles to each other (Reymanowna 1963, text-fig. 8A.)
The stomata of P. heterophylla appear to be more heavily cutinized than in other
species. For instance in P. ephela and P. grojecensis the guard cells are heavily
cutinized at the outer edges and along the stomatal aperture but in P. heterophylla
the whole upper surface of the guard cells is heavily cutinized, but sometimes slightly
thinner around the aperture.
The polar appendages of P. heterophylla are unique among the species under con-
sideration for the rest have thin polar areas.
Papillate subsidiary cells like those in P. heterophylla are also present in P. ephela,
P. minuta and P. ensiformis while P. grojecensis has a cuticular diaphragm on the
surface of the guard cells around the aperture.
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ENGLISH WEALDEN FLORA I 253
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EXPLANATION OF PLATES
Most of the figured specimen, are in the British Museum (Natural History) and bear the
prefix V.
GEOL. 17, 5
PLATE i
Circonitella knowltoni (Seward)
FIGS, i, 2. Gyrogonites showing five spiral cells meeting at the apical pole. Fig. i, V. 5 1552,
Fig 2, specimen lost. X 80.
FIG. 3. Basal pole of gyrogonite showing circular basal plate. V. 51553. x 80.
FIG. 4. Pyritised, unsquashed gyrogonite with convex spiral cells, showing basal plate.
V. 51554. x 80.
FIG. 5. Neotype with convex spiral partly broken away. V. 5 1555. x 80.
FIGS. 1-5, individual specimens all from one block numbered V. 5 1548 (re-registered from
V.i 0700).
Pachypteris lanceolate Brongniart
FIG. 6. Apical part of pinna, lower side uppermost, showing prominent midrib. "^3245.
X 2.
Bull. Br. Mus. mat. Hist. (Geol.) 17, 5
PLATE i
GEOL. 17, 5
PLATE 2
Hepaticites zeilleri (Seward)
FIG. i. General form of thallus showing dichotomous branching. Holotype, V. 233005 x i.
FIG. 2. Thallus showing thick midrib region and delicate lamina also rhizoids including
detached bunches. V.233oa. x 3.
FIG. 3. Enlargement showing rhizoids pointing forwards between a dichotomy of the thallus.
V.233oa. x 10.
Hepaticites ruff or di sp. nov.
FIGS. 4, 5. Thallus showing arcuate grooves, Fig. 4 immersed in paraffin, Fig. 5 coated with
ammonium chloride. Holotype, V.2343. x 3.
Becklesia sulcata sp. nov
FIG. 6. Part of a pinna showing stomatal grooves. V. 5 1540. x 20.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 5
PLATE 2
I
PLATE 3
Hausmannia dichotorna Dunker
FIG. i. Small fan-shaped leaf with broken apices. Sterile. ¥.12317. x 3.
FIG. 2. Larger more divided leaf with strap-shaped segments. Sterile. ¥.12349. X 3.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 5
PLATE 3
PLATE 4
Ruffordia goepperti (Dunker)
FIG. i. Frond with narrowest form of segments. V.2I57. x i.
FIG. 2. Frond with wider sterile pinnules at the top. Fertile pinnae at the base scarcely
visible (photographed dry) . V.2295. x I-
FIGS. 3, 4. Fronds with widest pinnules. Fig. 3, V.2357. xi. Fig. 4, ¥.12331. xi.
FIG. 5. Partof V.2I57. x 3.
FIG. 6. Sterilepart of V.2295. x 3-
FIG. 7. Pinnaof V.2357 showing ventation. x 3-
Bull. Br. Mus. nat. Hist. (Geol.) 17, 5
PLATE 4
PLATE 5
Ruffordia goepperti (Bunker)
FIG. i. Specimen with sterile top and fertile base, immersed in paraffin. V.2295. x I
FIG. 2. Fertile frond. V.2i6o. x i.
FIG. 3. Fertile frond. V.2i92«. x i.
FIG. 4. Frond in Fig. 3 enlarged to show the undulating margins of the fertile pinnules.
V. 21920. x 3.
Bull. Br. Mus. nat. Hist. (Geol). 17, 5
PLATE 5
PLATE 6
Pelletieria valdensis Seward
FIGS, i, 2. Largest fertile specimen picked out of the matrix. V.23&8. Fig. i, x i, Fig.
2, X 3.
Aspidistes sewardi sp. nov.
FIG. 3. Specimen immersed in paraffin, sori visible on most of pinnules. Holotype, V. 21730
x i.
FIG. 4. Middle part of same specimen enlarged to show rachis, sori and rolled edges, x 3.
Weichselia reticulata (Stokes & Webb)
FIG. 5. Part of pinna showing typical form with fleshy pinnules. Indication of reticulate
venation just visible in two bottom-left pinnules. V. 21740. X 3.
Pseudotorellia heterophylla sp. nov.
FIG. 6. Cuticle of upper surface showing cells in " packets ", Geological Survey & Museum
PF2758, x 200.
FIG. 7. Cuticle of upper surface from same specimen showing sculptured surface of the
cells, x 500.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 5
PLATE 6
PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING
TWO NEW DICYNODONTS FROM
THE TRIASSIC NTAWERE
FORMATION, ZAMBIA
C. B. COX
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 6
LONDON: 1969
TWO NEW DICYNODONTS FROM THE
TRIASSIC NTAWERE FORMATION, ZAMBIA
BY
C. B. COX
Department of Zoology, King's College, London
Pp. 255-294; 23 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 6
LONDON: 1969
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 17, No. 7 of the Geological
Palaeontological series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation :
Bull. Br. Mus. nat. Hist. (Geol.).
Trustees of the British Museum (Natural History) 1969
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 7 March, 1969 Price Seventeen Shillings
I
TWO NEW DICYNODONTS FROM THE
TRIASSIC NTAWERE FORMATION, ZAMBIA
By CHRISTOPHER BARRY COX
CONTENTS
Page
I. HISTORICAL INTRODUCTION ....... 257
II. STRATIGRAPHY ......... 258
III. SYSTEMATIC DESCRIPTIONS ....... 260
Genus Zambiasaurus submersus nov. .... 262
Genus Sangusaiirus edentatus nov. ..... 286
IV. AGE OF THE NTAWERE FAUNAS ...... 290
V. ACKNOWLEDGMENTS ........ 293
VI. REFERENCES ......... 293
SYNOPSIS
Two new types of dicynodont from the upper fossiliferous horizon of the Ntawere Formation
of Zambia are described. The first, Zambiasaurus submersus, is represented by the fragmentary
remains of at least eighteen juveniles and one adult, all of which were apparently drowned in a
sudden flood. Zambiasaurus is a stahleckeriid closely related to, and probably directly ancestral
to, Stahleckeria of the Middle Triassic of Brazil. It is the earliest known stahleckeriid, and the
first known outside South America. The earlier history of the group is unknown.
The other new dicynodont, Sangusaurus edentatus, is known only from a few fragments of the
skull. It is a kannemeyeriid, and shows some similarities to 1 schigualastia of the Middle or
Upper Triassic of Argentina.
Both the faunas of the Ntawere Formation appear to be intermediate in age between the
Cynognathus Zone fauna of South Africa and the Manda fauna of East Africa. Its age, in terms
of the standard Triassic sequences, is provisionally estimated as lowermost middle Triassic
(Lower Anisian).
I. HISTORICAL INTRODUCTION
THE Luangwa River is a major tributary of the Zambezi River, and is the main
river of the eastern half of Zambia. Fossil reptiles were first found in the upper
Luangwa River valley by Prentice in 1925, and the area was reconnoitred and later
investigated by Dixey in 1928 and 1935 (see Dixey 1936, 1937). He recorded fossil
reptiles from six different horizons within the Karroo succession, and also from a
higher horizon which he named the " Dinosaur Beds " and which he regarded as
Cretaceous in age.
The upper Luangwa River valley is remote, and is inaccessible to vehicles unless
these can negotiate rough dirt tracks. As a result, the area was not re-examined
until 1960 and 1961, when Dr. A. R. Drysdall of the Geological Survey of Northern
Rhodesia, and James Kitching, Field Officer of the Bernard Price Institute for
Palaeontological Research of Witwatersrand University, spent a total of four months
in the area. They examined the area north-west of the small village of Sit we, on
the River Luwumbu, 125 miles north of Lundazi, the administrative centre of this
north-eastern portion of the Eastern Province of Zambia. They confirmed that the
area was rich in fossil reptiles, and collected nearly 500 specimens.
GEOL. iy, 6. 23
258 TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
Karroo fossils were, of course, originally found in South Africa, and many of the
early specimens were sent to the British Museum (Natural History) in London for
appraisal and description. The resulting collection is of historic interest and contains
many type specimens, but many are imperfect and most of them are poorly docu-
mented and lack post-cranial elements. It was therefore decided to mount a major
expedition to Africa, with the aim of making a large and representative collection
of fossil Permian and Triassic vertebrates, using up-to-date methods of collecting
and recording full details as to their localities and stratigraphical origin. In view
of the known richness of the area, it was decided to rely upon the upper Luangwa
River valley for the collection of the bulk of the Permian fossils, and afterwards to
visit the south-western region of Tanganyika, where fossil vertebrates of both
Permian and Triassic age were known to occur.
The resulting expedition, the British Museum (Natural History) and University
of London Joint Palaeontological Expedition to Northern Rhodesia and Tanganyika,
spent six weeks collecting in the Sit we area in the summer of 1963, and a preliminary
account of the results has been published (Attridge, Ball, Charig & Cox 1964).
The expedition was greatly helped by the presence of James Kitching, whose services
had been very kindly loaned by the Bernard Price Institute. As a result there was
no difficulty in finding the fossil localities, once the necessary tracks for vehicle
access had been constructed, and a total of 220 specimens, weighing some 2\ tons,
was collected.
II. STRATIGRAPHY
As a result of their examination in 1960-61, Drysdall & Kitching were able to
give a detailed account of the stratigraphy and geology of the area (Drysdall &
Kitching 1963), from which the details in this section are taken.
Some alterations to Dixey's earlier interpretation of the stratigraphy of the area
became necessary. Drysdall & Kitching found that all of Dixey's six Karroo
horizons were part of a single richly fossiliferous formation, which they named the
Madumabisa Mudstone. The fauna of this is identical with that of the Lower
Beaufort Endothiodon and Kistecephalus zones of South Africa.
Above the Madumabisa Mudstone, and separated from it by an hiatus of probably
minor nature, Drysdall & Kitching defined a major lithological unit which they
have subdivided to give three formations of more convenient thickness. The lowest,
the Escarpment Grit, is of varying thickness (230- over 700 feet), whilst the upper-
most, the Red Marl, is 230-400 feet thick. Between the two lies the Ntawere
Formation, again of varying thickness (400- over 3,500 feet) and consisting of a
succession of thin, alternating arenaceous and argillaceous beds. The whole unit
is subdivided into formations on the basis of the different proportions of mudstone
and grit.
Above the Red Marl lies the non-fossiliferous Upper Grit, over 3,000 feet thick;
in some places the two are separated by a thin transition zone of intercalated grits
and mudstones, while in others there is a sharp transition.
Drysdall & Kitching found two fossiliferous horizons in the Ntawere Formation
and Red Marl. The lower horizon lies within the Ntawere Formation; its fauna
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 259
consisted predominantly of large dicynodonts, with fairly common large amphibians,
and also yielded an excellent Diademodon skull which has already been described
(Brink, 1963). The upper horizon comprised the uppermost beds of the Ntawere
Formation and the lower part of the Red Marl; its fauna contained the remains of
both vertebrates and molluscs.
Drysdall & Kitching consider that Dixey's " Dinosaur Beds " can only be identi-
fied, both lithologically and geographically, with the fossiliferous Ntawere Forma-
tion. Dixey based his assessment of the " Dinosaur Beds " upon some large, frag-
mentary, rolled bones which he found in situ and which Swinton tentatively
identified as possibly dinosaurian. However, Drysdall & Kitching found no bones
in situ at the localities indicated on Dixey's map, but they were able to identify a
thin superficial pebble deposit which Dixey regarded as the last debris of the almost
completely eroded Dinosaur Beds. This pebble deposit contained large fragmentary
bones, mostly identifiable as dicynodont. Bones of comparable size and appearance
are common in the nearby Madumabisa Mudstone, so that the bones found in the
pebble deposit are probably derived fossils originating in the Madumabisa Mudstone.
The bones observed in situ by Dixey have not been relocated and may also have
been derived fossils. Alternatively they may have come from as yet unrecognized
fossiliferous beds of the Ntawere Formation, which is now known to contain pseudo-
suchians — whose remains are not readily distinguishable from those of dinosaurs.
The fossiliferous beds themselves are dark red, soft mudstones, with feldspathic
grit bands. Bones almost always occur individually; no articulated remains have
been found. The bones are usually cracked and weathered, suggesting pro-
longed exposure before burial; they are often covered with a purple-red layer of
haematite, or with a thick layer of calcite. Fragments of bone are common in
some of the grit bands, and may be the remains of skeletons disarticulated by
shortlived floods which were responsible for the coarser sediment forming the grit
bands.
Drysdall & Kitching state (1963 : 22) that amphibians and molluscs (Unio
karooensis] are present towards the base of the upper fossiliferous horizon, where the
commonest reptiles are the cynodonts. The cynodonts persist into the higher
levels, where the predominant reptile is now a large dicynodont, where the reptiles
outnumber the amphibians, and where the molluscs are now rarer. One of the
cynodonts has already been described (Brink, 1963) as Luangwa drysdalli.
The fossils are found in the systems of small gulleys and eroding flats at the upper
ends of the small tributaries of the River Sangu. Each of the dicynodont genera
described in this paper was found in an individual collecting area of this kind, within
the general area described as " locality 15 " by Drysdall & Kitching, and hence within
the upper fossiliferous horizon. During the 1963 expedition we were not able to
define the stratigraphic inter-relationships of these individual areas, so that the
positions, within this horizon, of the two dicynodont genera described here are not
known.
Fossils from the Ntawere Formation are usually fairly scarce and covered with
layers of haematite or calcite. The Zambiasaurus material makes an outstanding
exception to this rule: a portion, about ten yards long and seven yards wide, of the
ENTOM. 17, 7. 23§
260 TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
side of a small ridge was found to be strewn with fragments of bone, nearly all of which
were completely free from matrix. The whole area was stripped of vegetation and
swept clean, with the result that 500-600 fragments of bone were collected. Though no
bones could be found in situ, some of the fragments were embedded in pieces of
coarse, unsorted, highly feldspathic grit. The proportion of the alkali feldspar, quartz
and mafic minerals suggests that the parent rock was of granite composition. The
poor sorting and lack of rounding of the grains, and unaltered condition of the
feldspar, suggests rapid deposition. The whole collection was therefore deposited
by one of the floods mentioned above.
III. SYSTEMATIC DESCRIPTIONS
Genus ZAMBIASAURUS nov.
The new genus has been named Zambiasaurus after the new name of its country
of origin.
GENERIC DIAGNOSIS : Large dicynodont : the composite restored immature skull
is 23 cm. long and 21 cm. broad, while the adult might have had a skull about 45 cm.
long and 40 cm. broad. No teeth in upper or lower jaws. Greatest width of skull
is across occiput; skull tapers anteriorly. Wide inter-orbital region, narrow inter-
temporal region. Blunt snout. Short median suture between nasals. Preparietal
bone absent; pineal foramen completely surrounded by parietals. Parietals are
slightly concave antero-posteriorly and form bulk of intertemporal bar. No sharp
median intertemporal ridge. Interparietal bone does not extend far forwards.
Sharp transition between dorsal and occipital surfaces. Occipital wings of squa-
mosal extend laterally and somewhat posteriorly. Palatal surface of premaxilla
bears pair of anterior ridges.
At least four sacral ribs. Scapular blade tall and narrow, with low ridge running
up antero-external edge; acromion process probably poorly developed. Coracoid
foramen wholly within precoracoid bone. Small pubis.
TYPE SPECIES: Zambiasaurus submersus sp. nov.
MATERIAL: Unless otherwise stated, all specimen numbers refer to the collection
deposited in the British Museum (Natural History).
The bone fragments were mainly parts of the post-cranial skeleton. Apart from
unidentified fragments and portions of ribs, the collection included :
Vertebral centra . . 58 Ulna .... 29
Sacral ribs ... 7 Ilium .... 40
Scapula .... 39 Ischium 16
Coracoid .... 13 Pubis .... 9
Precoracoid ... 4 Femur . . . . 51
Clavicle .... 6 Tibia . . . .24
Sternum .... 3 Fibula .... 13
Interclavicle . . . Metapodials, podials etc. . 15
Humerus .... 68 Premaxillae ... 4
Radius .... 19 Maxillae .... 9
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 261
Nasals .... 2 Quadrates .... 2
Frontals .... 6 Squamosals . . . 27
Postorbital . . . I Dentaries .... 2
Intertemporal region . . 8 Articular region of lower jaw 10
Braincase elements . . 8 Other fragments of lower jaw 6
As found, none of the limb bones was complete, but it was subsequently found that
the fragments included the whole of an ulna, two femora and two tibiae. Apart
from a few cynodont bones, and one other exception discussed below, there is no
variation in the morphology of the bones, which therefore appear all to belong to a
single species of dicynodont. This species is represented by the remains of at least
eighteen individuals, as there are eighteen right distal ends of humeri; however, it
is almost certain that there was really a somewhat greater number of individuals,
all of which are only incompletely represented. These individuals come from a
fairly restricted size range; in the humeri, for example, the proximal ends are from
8-5 to io-o cm. across and the distal ends are from 7-8 to n-o cm. across.
In addition to these individuals, there are the remains of a single individual of
considerably larger size, the distal end of whose humerus is 17 cm. across. This
specimen could be regarded either as a larger member of the same species as the
numerous smaller individuals, or as belonging to a different, larger species.
There are several reasons for believing that the many smaller individuals are
immature forms, and that the single large individual represents the adult of the
same species. The immaturity of the small specimens is indicated by the lack of
co-ossification and sutural union of the skull bones (even the bones of the braincase
have not become firmly united) and by the poorly defined nature of the articular
surfaces of the limb bones. On the humerus, for example, the articular areas for
the glenoid, radius and ulna are hardly defined at all, while the articular surfaces
of the radius, tibia and fibula are almost featureless and give the impression of
having been covered in life with a thick capping of cartilage. Such surfaces on this
and other bones have a characteristic appearance: the surface is smooth but inter-
rupted by a large number of tiny round holes, the edges of which are often slightly
raised, giving the appearance of tiny volcanic craters. In living reptiles this appear-
ance is characteristic of surfaces covered by cartilage, and it will be referred to as
" cartilage ornament " in the descriptive sections below.
The bones of the larger individual, on the other hand, have very well developed
articular areas, but are otherwise identical with those of the smaller individuals.
It is therefore regarded as the adult of the species, but is described separately, after
the smaller individuals have been described, so as to facilitate any taxonomic change
in its status, should this subsequently be found necessary.
A death-assemblage composed of a single adult and many juveniles is rather
unusual. The flood which deposited the accompanying coarse grit may provide a
possible explanation ; it is conceivable that such a flood might cause a heavy mort-
ality in the young dicynodonts, but that only a few of the larger adults would be
caught and overwhelmed. The adult might even be a carcass of an individual
which had died recently and which was caught up by the flood waters.
262 TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
Zambiasaurus submersus sp. nov.
Text-figs. 1-20
The specific name submersus refers to the likelihood that the specimens were
drowned.
HOLOTYPE OF Z. submersus: Livingstone Museum (Zambia) specimen No.
LM/NH 9/2, consisting of interparietal, left parietal and part of left squamosal of
presumed immature individual.
PARATYPES: British Museum (Natural History) catalogue Nos. R. 9001^.9140;
Livingstone Museum (Zambia) specimen Nos. LM/NH 9/3-9/35.
HORIZON AND LOCALITY : Upper fossilif erous horizon of Triassic Ntawere Formation ;
from locality 15 of Drysdall & Kitching (1963), about 3^ miles west of Sitwe, in the
upper Luangwa Valley, Eastern Province, Republic of Zambia (Field No. 15 A/I).
DESCRIPTION. As explained below the original skull morphology has had to be
reconstructed and deduced from the eighty-five fragments listed above.
The most extensive fragment, which has been designated as the type specimen
(Livingstone Museum specimen no. LM/NH 9/2; cast in B.M.(N.H.) is R.gooo)
comprises an interparietal, left parietal and part of the left squamosal (Text-fig,
i, b-e). The right parietal has become detached and lost, so that the sutural
surfaces by which it was attached to the interparietal and left parietal are visible.
It can therefore been seen (Text-fig, id) that the interparietal does not extend far
forwards between the two parietals. The interparietal also does not extend far
laterally before it joins the squamosal, neither does it overlap the occipital surface
of the squamosal to any great extent. Though there is a sharp transition between
the dorsal surface of the intertemporal bar and the posterior surface of the inter-
parietal, there is also a recessed area in the dorso-median region of the interparietal,
where the nuchal ligaments were presumably attached. Below this recess lies the
occipital surface of the interparietal, in which there is a pair of nutrient foramina.
The dorsal surface of the left parietal is smooth and slightly concave antero-
posteriorly. Though it is also slightly concave transversely, so that there would
have been a very slight median ridge, the intertemporal bar as a whole is flattened,
rather than forming an inverted V shape. The dorsal and lateral surfaces of the
parietal meet at an acute angle (about 60°) and the lateral surface is shallowly
concave dorso-ventrally. The posterior end of the lateral surface of the parietal
is slightly damaged, but extended back to overlap the squamosal. The upper part of
the lateral surface of the parietal is slightly recessed and bears the pitted and grooved
ornament characteristic of a surface overlapped by another bone. There can be
no doubt that this surface was covered by the posterior extension of the post-
orbital. Another specimen (R . 9020) shows this surface even more plainly, and it
is clear that the postorbital extended back as far as the junction between the parietal
and the interparietal.
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
263
The parietal in the holotype forms the whole of the border of the left half
of the pineal foramen (Text-fig, ib). Another specimen (R.9020) in which this
region is slightly better preserved, shows that the parietal forms a thin strip of the
dorsal surface both lateral to, and anterior to, the pineal foramen (Text-fig. la).
In neither specimen is there any trace of a suture delimiting a separate preparietal
bone, and this bone must have been absent.
D
VSUf.
FIG. i. Zambiasaurus submersus gen. et sp. nov., juvenile, x f . A, dorsal view of anterior
end of left parietal, R.go2o. B-E, holotype (Livingstone Museum No. LM/NH 9/2)
in B, dorsal view; C, lateral view; D, medial view; E, anterior view. Abbreviations:
d.su.f., dorsal sutural area for frontal; IP, interparietal ; P, parietal; pin., pineal opening;
rec.pf?, recess for Ppostfrontal; SQ, squamosal; su.p., sutural area for right parietal;
su.po., sutural area for postorbital; v.su.f., ventral sutural area for frontal.
264
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
Much of the anterior end of the dorsal surface of the parietal is covered by ridged,
grooved and pitted areas of overlap (Text-fig. ia, b). The most medial of these
areas must have received the posterior end of the frontal (Text-fig, ia, b) which
therefore reached as far posteriorly as the level of the posterior border of the pineal
foramen and also approached very close to its lateral and anterior borders (Text-
figs. ia, 2b, 2ob). The most lateral area of sutural overlap on the antero-dorsal
region of the parietal is that for the postorbital. Between this and the area for the
frontal there is a deep posteriorly-directed recess (Text-fig. la-c: rec.pf ?). This
N
FIG. 2. Zambiasaurus submersus gen. et sp. nov. Drawings showing relationships of
preserved fragments of skull to reconstruction of complete juvenile skull, x J. A, lateral
view; B, dorsal view; C, occipital view. Abbreviations: F, frontal; N, nasal; MX,
maxilla; PMX, premaxilla; Q, quadrate; SQ, squamosal.
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 265
recess may have received a small separate postfrontal bone. If it was present, this
bone may have been concealed by a superficial meeting between the postorbital and
the frontal.
There is a considerable angle between the more posterior part of the parietal and
those areas to which the frontal attached. This fact, and the concave outline of the
dorsal edge of the parietal in lateral view (Text-fig, ic), suggests that the intertemporal
bar projected postero-dorsally above the level of the rest of the dorsal surface of
the skull (Text-fig. 2a). This is confirmed by fragment R.goi4 (Text-fig. 3a), which
comprises the region around the front end of the pineal foramen. The line of the
dorsal edge of the intertemporal bar apparently continued forwards for a short
distance on to that part of the frontal which lies antero-lateral to the pineal foramen,
for this part of the bone is slightly thickened above the level of the rest of the frontal.
This fragment also confirms that there is no preparietal bone and that the frontal
overlaps the anterior end of the parietal. The frontal also extends posteriorly for a
short distance under the parietal, where there is a wide sutural union between these
two bones (Text-fig, id, e: v.su.f.).
The morphology of the area immediately in front of the pineal foramen, including
the interorbital width, is shown by specimen R.goiS. This is a large piece of a right
frontal, including both the midline suture and part of the edge of the orbit. Speci-
men R .9016 is an almost identical fragment of a left frontal, on the postero-medial
region of which can be seen the suture for attachment to the parietal. This fact
is important, for it establishes the relationship between these frontal fragments, inclu-
ding the upper margin of the orbit, and the intertemporal bar. The resulting position
of the orbit is relatively far back, so that the posterior edge of the post-orbital bar
is at the level of the pineal foramen.
It is possible, but not certain, that the most antero-lateral corner of the frontal
fragment R . 9015 bears a short stretch of the surface to which the prefrontal bone was
attached. Even if it does not, the suture cannot have been far anterior to this point,
since that bone normally forms the antero-dorsal corner of the orbit.
Specimen R.90I2 is part of a right nasal bone (Text-fig. 3b, c). The bone was
overlapped postero-medially by the anterior end of the frontal, and antero-medially
by the posterior end of the premaxilla. These surfaces are separated by only a
short median sutural surface for the left nasal. In another, slightly larger, right
nasal fragment (specimen R.90I3) the overlap surfaces for the premaxilla and frontal
meet, completely covering the median suture between the two nasals. The antero-
lateral corner of these nasals bears a number of foramina.
The dorsal surface of the nasal R.90I2 consists of two planes, which meet along a
line which runs antero-laterally. The more postero-lateral plane faces dorso-
laterally and represents the lateral surface of the snout. The anterior plane faces
antero-dorsally and represents the beginning of the down-turning of the snout.
This plane provides a clue as to the relative position of the nasal and of the anterior
end of the snout since, together with the anterior surface of the premaxilla, it must
form a smooth curve when seen in lateral view (Text-fig. 2a). Some hint as to the
distance between the nasal and the anterior end of the premaxilla is also provided by
the angle of the suture between these two bones, as seen along the front surface of
266
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
the nasal. In most dicynodonts this suture runs to the antero-dorsal corner of the
nostril in an almost straight line.
The front edge of the premaxilla is blunt, running transverse to the main
axis of the skull. The grooves which lie lateral to the paired anterior palatal
ridges therefore run on to the anterior edge of the premaxilla (specimen R.QOOI;
Text-fig. 3d).
pin.
su.n.
su.m.
FIG. 3. Zambiasaurus submersus gen. et sp. nov., juvenile, x f . Dorsal view of fragment
R.QOI4 comprising area around anterior end of pineal foramen. B, C, fragment of right
nasal (R.goia) in dorsal (B) and medial (C) views; D, ventral view of premaxilla (R.gooi).
Abbreviations: F, frontal; P, parietal; pin., pineal foramen; su.f., sutural area for frontal;
su.m., sutural area for maxilla; su.n., sutural area for left nasal; su.pmx., sutural area for
premaxilla.
There is little difficulty in fitting a maxilla (e.g. specimen R . 9004) to the premaxilla.
All nine maxillae are tuskless. Like the premaxilla and the anterior part of the
nasal, the maxilla bears a number of nutrient foramina. There is a slight posteriorly
directed flange down its postero-lateral edge.
At no point is there any junction between the preserved parts of the nasal-
premaxilla-maxilla section and the more posterior section of the skull. It was
therefore only possible to attempt to estimate the original relationship between
these two sections by assembling each independently in a plasticene matrix and then
matching them together, keeping the palatal surface of the premaxilla horizontal.
The reconstruction shown appears quite plausible and does not violate any known
normal feature of dicynodont morphology. Nevertheless, the exact distance between
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 267
the two sections (and therefore the exact length of the frontal) must remain in
doubt.
At the other end of the skull, there is similarly no certain relationship between the
intertemporal-interparietal region and the remainder of the squamosal and braincase
(Text-fig. 2). Most of the fragments of squamosal comprise the region where the
zygomatic arch arises from the front surface of the occipital wing. The orientation
of the base of the zygomatic arch, and also the orientation of the piece of squamosal
attached to the interparietal of the type specimen (Text-fig, ib) show that the
squamosals extended laterally and somewhat posteriorly from the midline. The
approximate width across the occiput follows from the interorbital width, since the
zygomatic arches must have continued anteriorly and somewhat medially into the
suborbital bar.
The fragments of braincase include a good basioccipital-basisphenoid (specimen
R . 9027) and a good exoccipital-opisthotic-prootic (specimen R . 9028) . The occipital
condyle is of the normal tripartite dicynodont pattern. The tuber around the
fenestra ovalis is formed partly by the basisphenoid and partly by the opisthotic.
The opisthotic also forms the whole of the distal end of the paroccipital process;
its anterior surface is covered by the prootic. Dorso-laterally, both the prootic
and the opisthotic end abruptly in a thick surface, the ornament of which shows that
it was capped by cartilage, and which presumably met the squamosal. The dorsal
edge of the opisthotic, which slopes upwards and outwards, bears a similar ornament
and must have joined the supraoccipital. The medial surface of the prootic and
opisthotic bears the excavation for the inner ear.
The quadrate is of usual dicynodont type. Once again, its exact relationship to
the remainder of the skull is uncertain, and it has merely been placed in the position
normal in the group, at about the same level as the ventral margin of the premaxilla
and ventro-lateral to the distal end of the paroccipital process.
Most of the fragments of lower jaw are of the articular region, which is of normal
dicynodont type. Two fragments of the front end of the dentary (specimens
R.9039, 9040) show that this was wide and blunt, as might have been expected from
the shape of the premaxilla.
Postcranial morphology. To facilitate comparison, all bones are illustrated as
viewed from the left side. Where necessary, bones from the right side have been
reversed in the drawings so as to appear as left side bones.
There is a considerable number of fragmentary vertebrae. The prezygapophyses
are usually concave, the postzygapophyses convex. The antero-dorsal corner of
the lateral surface of the centrum bears a small facet for the lower end of the rib
head. The remainder of the facet for the rib head extends postero-dorsally up the
side of the neural arch. This facet extends to a varying degree up the ventral
surface of the transverse process, which is dorso-laterally directed.
It is not possible to obtain any useful information from the fragments of ribs.
The scapula illustrated (Text-fig. 4) is reconstructed from two complementary
fragments: a lower fragment (R.goGS) and an upper fragment ^.9069). The blade
is very long and narrow. There is a rather low scapular spine which runs for some
distance up the antero-lateral edge of the bone (Text-fig. 4b) but which does not
GEOL. 17, 7.
268
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
FIG. 4. Zambiasaurus submersus gen. et sp. nov., juvenile. Composite restoration. A, lateral
view of scapula, precoracoid and coracoid (scapula and precoracoid are composite recon-
structions) ; B, section through scapula at level a-b ; C, view of ventral end of scapula, lateral
surface uppermost, x f. Abbreviations: cor.f., coracoid foramen; rug. trie., rugosity
for ligament of triceps muscle.
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
269
reach its upper end. The acromion process is damaged, but does not appear to
have been very large. There is a pronounced rugose area on the postero-dorsal
edge of the scapula a short distance above the glenoid; this is probably the area of
FIG. 5. Zambiasaurus submersus gen. et sp. nov., juvenile. Coracoid in A, dorsal view
and B, medial view, x f
FIG. 6. Zambiasaurus submersus gen. et sp. nov., juvenile. Sternum in dorsal view, x f.
origin of the scapular ligament of the triceps muscle. The glenoid surface is sub-
circular, but the surface for attachment of the precoracoid is quite thin (Text-fig. 40) .
The coracoid illustrated (specimen 1^.9073; Text-figs. 4, 5) is probably from a
somewhat smaller individual than the scapula illustrated, as the scapular facet
on the coracoid is narrower than the coracoid facet on the scapula. The outlines
of these two surfaces do not, in any case, match; this is probably because there was
270
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
still much cartilage between the elements of the girdles of these presumably juvenile
individuals.
The precoracoid (Text-fig. 4) is incomplete; it is restored from three incomplete
specimens (R. 9078-80). Part of the edge of the coracoid foramen is preserved;
since there is no notch in the lower edge of the scapula, this foramen must have
lain wholly within the precoracoid.
Though a few fragments of clavicle are preserved, these give no useful information
about the bone. No fragments of interclavicle have been identified.
The sternum (Text-fig. 6) is roughly hexagonal in outline. Its ventral surface is
slightly concave. On its dorsal surface lie a pair of postero-lateral bosses, on the
ends of which can be seen cartilage-ornament. These bosses probably gave insertion
to the ventral ends of the anterior thoracic ribs.
Since no complete humerus is known, it has reconstructed mainly from two comple-
mentary fragments, a proximal fragment R.goSS and a distal fragment
A B
FIG. 7. Zambiasaurus submersus gen. et sp. nov., juvenile. Composite restoration of
humerus in A, ventral view and B, posterior view, x f . Abbreviations : dp.crest, delto-
pectoral crest; ent.f., entepicondylar foramen.
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 271
(Text-figs. 7, 8). It is strongly twisted. As in all the limb-bones, the areas of
muscular insertion and of articulation are poorly defined, as might be expected in
juvenile animals. There is thus little trace of the proximal condyle, nor of the
condyles for the radius and ulna. Traces of cartilage-ornament can be seen over
many of these surfaces, proving that the absence of the condyles is not due simply
to erosion .
No complete radius is known, and the bone has been reconstructed (Text-fig. 9)
from two proximal fragments (specimens 1^.9093, 9094) and two distal fragments
(specimens R.gogs, 9096). It is a slender bone and has been reconstructed to be,
as is normal, slightly longer than the ulna. The proximal end is slightly convex,
the distal end slightly concave.
A single complete right ulna is known (specimen R.gog8, Text-fig. 9). The bone
is fairly slender. Only the horizontal surface of the glenoid notch is represented.
Since the whole of the proximal surface of the ulna is preserved (as proved by its
covering of cartilage-ornament) , the remainder of the sigmoid notch must have been
B
FIG. 8. Zambiasaurus submersus gen. et sp. nov., juvenile. Composite restoration of
humerus in A, dorsal view and B, anterior view, x f. Abbreviations: dp.crest,
delto-pectoral crest.
272
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
borne on a separately-ossified olecranon process. The distal end of the ulna is
strongly convex.
Like those of the pectoral girdle, the bones of the pelvic girdle (Text-figs. 10-12)
cannot be fitted together accurately, and much of the regions between the bones
must still have been cartilaginous at this stage of growth.
The outline of the ilium has been based mainly on that of specimen R. 9103, except
for the extreme posterior preserved portion of the blade which is taken from speci-
men R.9io6. The sacral facets are reconstructed from specimens R. 9103-05.
The outer surface of the blade is slightly concave both dorso-ventrally and antero-
posteriorly, while the inner surface is slightly convex in both these planes. The
A B
FIG. 9. Zambiasaurus submersus gen., et sp. nov., juvenile. A, radius (composite restoration)
and ulna in posterior and distal views. B, medial view of ulna; C, radius (composite
restoration) and ulna in anterior and proximal views, x f. Abbreviations: RD, radius;
UL, ulna.
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 273
acetabular surface of the ilium faces ventro-laterally and slightly anteriorly. The
facets for at least four sacral ribs can be distinguished on the inner surface of the
blade (Text-fig. 12).
Most of the outline of the ischium (Text-figs. 10, n) is based upon specimen R . 9108 ;
most of the upper edge and a little of the posterior edge is complete. Specimen
pu.tub.
FIG. 10. Zambiasaurus submersus gen. et sp. nov., juvenile. Lateral view of pelvis. X
Abbreviations: obt.l, obturator foramen; pu.tub., pubic tubercle.
274
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
IL
IS
FIG. ii. Zambiasaurus submersus gen. et sp. nov., juvenile. Views of acetabular-articu-
latory surfaces of the ilium (IL), ischium (IS) and pubis (PU). x f . Lateral surfaces of
the bones are peripheral, medial surfaces are central.
FIG. 12. Zambiasaurus submersus gen. et sp. nov., juvenile. Medial view of composite
restoration of ilium, x f .
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
275
R . 9109 includes a little more of the ventral portion of the bone. The anterior edge
of the ischium is notched for the obturator foramen. The postero-dorsal region
of the bone slants somewhat inwards, so that there is a stout ridge between this
surface and the more antero-ventral region.
All edges of the pubis (specimen R. 9113, Text-figs. 10, n) bear cartilage-ornament,
so that the figures show the bone complete at this stage of ossification. It bears a
large convex face which must include the surfaces which articulated with the ilium
and ischium, and also the pubic contribution to the acetabulum. Below this region
the pubis projects antero-laterally to form a rather elongate pubic tubercle. The
posterior edge of the bone is notched for the obturator foramen.
The femur illustrated (Text-fig. 13) is specimen R.girS, which is complete and
from the right side; it is i8'5 cm. long. The articular facets are poorly developed.
In side view it can be seen that the distal condyles were probably directed some-
what posterior to the main axis of the bone.
FIG. 13. Zambiasaurus submersus gen. et sp. nov., juvenile. Femur in A, anterior view ;
B, lateral view; C, posterior view, x f.
276
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
The tibia illustrated (Text-fig. 14) is specimen R.QI23, which is also complete and
from the right side. The proximal surface bears a pair of concavities for the femoral
condyles. The lateral surface of this end of the bone is also slightly notched where it
adjoined the proximal end of the fibula. The cnemial crest is represented by a
thickening of the antero-lateral surface, which ends proximally in a surface which
faces slightly anteriorly. The distal surface of the tibia can be divided into a
moderately flat lateral region, and a markedly convex medial region.
The fibula is a slender, slightly curved bone (Text-fig. 14). The proximal end
figured is specimen R . 9128, while the distal end is specimen R. 9129. The proximal
end is convex; it is crescentic in outline, so that its medial edge curves around the
lateral surface of the tibia. The distal articular surface is also convex, but is oval
in outline.
Though a number of elements from the hand and foot are preserved, no useful
information can be derived from them.
FIG. 14. Zambiasaurus submersus gen. et sp. nov., juvenile. A, tibia and fibula in posterior
and proximal views; B, medial view of tibia; C, tibia and fibula in anterior and distal
views, x f. Abbreviations: FIB, fibula; TIB, tibia.
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
277
ADULT SPECIMEN. Together with the remains of the immature specimens, there
was preserved also a number of much larger dicynodont bones. Since no individual
bone was duplicated, and since all of these bones were of commensurate size, it
seems likely that all are derived from a single individual. They have therefore all
been allocated a single specimen number, 1^.9140. They include a few uninformative
fragments of skull and ribs, a few neural arches, two vertebral centra, a complete
left coracoid and fragments of a pair of scapulae, a left humerus, a right radius,
ulna and tibia, and a single phalanx.
One of the neural arches is almost complete, and is shown (Text-fig. 15) mounted
upon one of the centra. The rib facet extends from the corner of the centrum up the
FIG. 15. Zambiasaurus submersus gen. et sp. nov., adult (R.gi4o). Neural arch and centrum
(not originally associated) in A, anterior view; B, lateral view; C, posterior view, x \.
side of the dorso-laterally directed transverse process, but does not reach its distal
end. The outline of the dorsal end of the neural spine is restored from another speci-
men from this large individual. It is rather narrow antero-posteriorly, and slopes
backwards at a considerable angle.
The outline and morphology of the scapula (Text-fig. 16) and coracoid are identical
to those of the immature specimen (cf. Text-fig. 4). In anterior or posterior view
it can be seen that the blade of the scapula is curved to conform to the outline of the
rib cage.
A fragment of the postero-dorsal corner of the proximal end of the humerus is
preserved and shows a well developed articular condyle (Text-fig. I7a). The distal
half of the bone (Text-fig, ryb, c) shows well developed condyles for the radius and
ulna.
278
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
\
\
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
279
The proximal ends of the radius and ulna, and the distal end of the ulna, are
preserved. This fragment of radius shows no features of interest. The olecranon
region of the ulna is lacking, and the proximal surface of the remainder of this end
of the bone is unfortunately too poorly preserved for it to be possible to ascertain
whether there was a separate olecranon ossification, as in other Triassic dicynodonts.
The proximal end of the tibia is well preserved (Text-figs. 18, 19). The pair of
depressions for the femoral condyles are clearly visible. The cnemial ridge running
up the antero-lateral face of the bone is better developed than in the immature
specimen (cf. Text-fig. 14), and terminates in a rounded surface which is directed
dorsally and anteriorly. A rugose area (Text-fig. 19, rug.) on the postero-lateral
FIG. 17. Zambiasaurus submersus gen. et sp. nov., adult (1^.9140). Humerus, x f.
A, B, dorsal views of fragments (A) of proximal end and (B) of distal end ; C, ventral view
of fragment of distal end. Abbreviation: prox. cond., proximal condyle.
a8o
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
corner of the bone may mark the position where the proximal end of the fibula
contacted the tibia.
If measurements (in cm.) of the bones of the adult are compared with those of the
immature specimens figured, the following results are obtained.
Length of scapula
Length of coracoid
Width of distal end of humerus
Width of proximal end of radius
Width of proximal end of tibia
Juvenile
24
6-3
9
4-6
Adult
43
10-5
17
TO
10-5
Adult/
Juvenile
1-8
1-7
1-9
2-2
2-1
cnem.
crest
B
FIG. 18. Zambiasaurus submersus gen. etsp. nov., adult. (R. 914°) Tibia, x £. A, medial
view; B, anterior and proximal views. Abbreviation: cnem. crest, cnemial crest.
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 281
It can be seen that the adult bones are about twice the size of the juvenile bones.
If the skull of the adult was, similarly, about twice the size of the restored juvenile
skull and retained a similar length : width ratio, it would have been about 45 cm.
long and 40 cm. wide across the occiput.
DISCUSSION. Taxonomy and relationships of Zambiasaurus. There can, first of
all, be little doubt that Zambiasaurus is a stahleckeriid, rather than a kannemeyeriid.
Its blunt snout, wide but low occiput, short temporal opening and lack of a
cnem.
crest
rug.-f*
FIG. 19. Zambiasaurus submersus gen. et sp. nov., adult (R. 9140). Tibia, x J. A, lateral
view; B, posterior view. Abreviation: cnem. crest, cnemial crest; rug., rugosity.
high intertemporal crest, are all similar to these diagnostic features of the family
Stahleckeriidae as originally defined (Cox, 1965).
Zambiasaurus is the first stahleckeriid known outside South America. The
Stahleckeriidae at present includes three genera: Stahleckeria of the Santa Maria
Formation of Brazil, described by von Huene (1935-42) ; Dinodontosaurus from the
above Brazilian Formation (Cox, 1965) and also from the Chanares Formation of
Argentina (Cox, 1968) and Chanaria, also from the latter Formation (Cox, 1968).
Both Dinodontosaurus and Chanaria have retained tusks and a preparietal bone,
in contrast to Stahleckeria in which both these features are absent. Their absence
in Zambiasaurus suggests that this new genus is more closely related to Stahleckeria
than to the other two stahleckeriid genera.
282
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
Though tusks are frequently lost in dicynodonts, the absence of the preparietal
is more rare. Comparison of the skulls of Zambiasaurus and Stahleckeria (Text-fig.
20) reveals several other similarities. Despite some superficial differences, the
morphology of the whole intertemporal bar is very similar in the two genera. The
parietals in Zambiasaurus can still be seen anterior to the pineal foramen, where they
have occupied the space normally filled by the preparietal. In Stahleckeria the
B
FIG. 20. Zambiasaurus submersus gen. et sp. nov., reconstructed juvenile skull x J (left)
and Stahleckeria potens, adult skull x c. 1/12 (right). A, lateral views: B, dorsal views;
C, occipital views. (Stahleckeria after Camp, 1956).
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 283
frontals have extended slightly further back, enclosing the anterior half of the pineal
foramen. In both genera the frontal meets the parietal at a level half way along the
pineal foramen, the parietal forms a rather wide intertemporal bar which is almost
flat in transverse section, and the interparietal contributes little to the intertemporal
bar.
Another similarity between Zambiasaurus and Stahleckeria is that in both the
nasals either meet for only a very short distance in the midline or are separated by
a junction between the frontals and the premaxilla. In Stahleckeria this is probably
a result of the very wide, massive snout ; if the nasals lie far apart, the processes of
ossification outwards from the centre of each bone might not reach the midline
before this area had been ossified by the frontal. As far as one can tell, the snout
of Zambiasaurus also was rather wide ; this follows from the shape of the incomplete
nasals, and the centre of ossification of the bone certainly lies more lateral than these
fragments.
As far as the post-cranial skeleton is concerned, it is in any case difficult to find
many systematic differences between the Triassic dicynodonts (see Cox, 1965 for a
general survey). This difficulty is compounded in Zambiasaurus by the fact that
most of the bones are known only in their juvenile state, with poorly developed
processes for muscular insertion and articular condyles. A few similarities between
Zambiasaurus and Stahleckeria can nevertheless be observed. The narrow, pos-
teriorly-directed neural spine of Zambiasaurus (Text-fig. 15) is very like that of the
anterior vertebrae of Stahleckeria. The coracoid, sternum and pelvis are in general
similar in the two genera, there being a striking similarity between the ischium and
pubis of Zambiasaurus and those of a juvenile Stahleckeria figured by von Huene
(1935-42, PI. 9, fig. 4). The difficulties mentioned above make it impossible to
make useful comparisons between the limb bones of the two genera.
The absence of tusks, and of the preparietal bone, the morphology of the inter-
temporal bar and of the snout, and some post-cranial features thus all suggest that
Zambiasaurus may be quite closely related to Stahleckeria. The Ntawere Formation,
in which Zambiasaurus is found, is certainly older than the Santa Maria Formation,
which contains Stahleckeria. It is therefore possible that Zambiasaurus may be
actually ancestral to Stahleckeria, and the known differences between the two genera
may now be examined with this possibility in mind.
As far as one can tell, the occipital wings of the squamosal are directed more
posteriorly in Zambiasaurus than in Stahleckeria, so that its occiput is not as flat
as that of Stahleckeria, and its temporal opening is slightly longer. The occiput
itself, as restored, is not as wide in Zambiasaurus as in Stahleckeria. However, this
feature could not, from the material preserved, be restored with any great degree of
accuracy. It is also worth noting that the restored skull is that of a juvenile and
that the occiput apparently does increase in relative size as the animal grows.
This is suggested by the relatively larger occiput of the large specimen of
Dinodontosaurus turpior which was described as a possible " old bull " (Cox, 1965,
pp. 489-494)-
Another difference between the occiputs of the two genera is that the interparietal
of Stahleckeria may extend further laterally, as shown in Camp's (1956) reconstruction
284 TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
based on further study of von Huene's material. However, the outline of this bone
is shown dotted in his figure (1956, Fig. 45) and this point is therefore uncertain.
In any case, there is no obvious reason why the interparietal should not become
wider during the evolution of the very wide occiput of Stahleckeria.
The snout of the " old bull " Dinodontosaurus mentioned above is also more massive
than that of the smaller specimens, and this should be borne in mind when comparing
the immature skull of Zambiasaurus with the adult skull of Stahleckeria, as these
show a similar difference. The snout of Stahleckeria is extremely heavy and short,
so that the centre of the maxilla lies under the anterior edge of the orbit. As
reconstructed, the maxilla of Zambiasaurus lies further forwards. However, the
exact inter-relationship between its maxilla and orbit cannot be determined from the
fragments available (cf. Text-fig. 2) and, in any case, the condition shown in Zambia-
saurus could well be ancestral to that found in Stahleckeria.
The interorbital region of Stahleckeria also differs somewhat from that of Zambia-
saurus in that its nasal and prefrontal extend further posteriorly, so that the anterior
half of the upper margin of the orbit is formed by the prefrontal instead of by the
frontal as in Zambiasaurus. This, too, may be a result of the shorter snout of
Stahleckeria, since the centres of ossification of the bones of this region must lie
further posteriorly, so that they will encroach on the area formerly occupied by the
frontal.
The only obvious difference between the post-cranial skeletons of Zambiasaurus
and Stahleckeria is the shape of the scapula. In Stahleckeria (von Huene, 1935-42,
PI. 7, figs. 1-3) the scapula narrows above the glenoid region, but further dorsally
it becomes progressively wider ; there is a strong, antero-laterally directed acromion
process, from which a stout spine runs up the outer surface of the blade near its
anterior edge. In Zambiasaurus, on the other hand, the scapula does not expand
dorsally, probably had a small anteriorly directed acromion process, and has a very
low spine. However, there is no reason to suppose that the above features of the
scapula of Stahleckeria could not have evolved from the condition found in
Zambiasaurus.
To summarize, none of the known differences between the two genera precludes
the possibility that Zambiasaurus is not only closely related to Stahleckeria, but is
also directly ancestral to it. It is impossible, on the basis of present material, to
test this hypothesis further ; such tests will depend on the discovery of stahleckeriids
of an age intermediate between that of the Ntawere Formation and that of the Santa
Maria Formation.
Origin of the stahleckeriids. As will be shown later in this paper, the age of
the Ntawere Formation may provisionally be estimated as lower Anisian. Zambia-
saurus is therefore the oldest known stahleckeriid. As has already been shown, it
is also probably ancestral to Stahleckeria and less closely related to the tusked genera
Dinodontosaurus and Chanaria. The line leading to these tusked stahleckeriids
must, then, have diverged from the tuskless Zambiasaurus-Stahleckeria line in or
before the Lower Triassic (Scythian).
I have earlier (1965) suggested that the stahleckeriids (including all the above
genera) may be separated from the kannemeyeriids on the basis of the form of the
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 285
palate and occiput. The morphology of these areas is, of course, determined by the
orientation and volume of the jaw muscles, and by the way in which the jaws are
used during feeding. The modifications in the kannemeyeriid line (seen most clearly
in Kannemeyeria and Ischigualastid) seem mainly to be directed towards increasing
the antero-posterior length of the jaw muscles. This has been achieved by some
exaggeration of that posterior extension of the squamosal which is common in
Permian dicynodonts, and also by the postero-dorsal extension of the posterior
region of the inter-temporal bar. These features seem to accompany a rather
pointed premaxilla (e.g. Sangusaurus, see Text-fig. 22b) in which the anterior ends
of the paired anterior palatal ridges of the premaxilla meet the converging lateral
walls of the mouth cavity. As a result of this, the grooves which lie lateral to these
ridges meet the antero-lateral margins of the snout, not its anterior edge.
The modifications of the stahleckeriids, instead, seem to have been directed
towards an increase in the width of the skull and anterior end of the jaws. The pos-
terior extension of the squamosal is lost altogether and the temporal opening is very
short, but that part of the squamosal which lies lateral to the root of the zygomatic
arch is greatly developed. This lateral development presumably provided attach-
ment for muscles which would have allowed more accurate control of the lateral
movements of the jaw than would have been possible in the kannemeyeriids. This
may be correlated with the transversely widened anterior surface of both the pre-
maxilla and the dentary. As a result of the width of this region, the paired palatal
ridges of the premaxilla, and the grooves lateral to these ridges, run on to the anterior
edge of the snout (Text-fig. 3d).
I have earlier (1965) suggested that these differences may be explained by their
feeding habits, the kannemeyeriids having been browsers feeding on fronds and leafy
branches, while the stahleckeriids were grazers, feeding on any low herb-cover similar
to that provided by grasses today. Further understanding of the functional mech-
anical differences between these two groups will be possible only when well preserved
undistorted skulls and jaws are available. These would permit the study of the
sequences of possible jaw movements, in the same way as Crompton & Hotton's
(1967) investigation of the functional morphology of two Permian dicynodonts,
Emydops and Lystrosaurus.
It is at present impossible to suggest confidently any links between the Triassic
dicynodont groups and the varied dicynodonts of the Upper Permian Kistecephalus
Zone. Immediately above this zone lies the Lystrosaurus Zone, in which tiny
Myosaums and the aberrant lystrosaurids are the only dicynodonts known. Above
this is the Cynognathus Zone from which Kannemeyeria is the only dicynodont
described. Though the Sinokannemeyeria fauna of China is about the same age
as the Cynognathus Zone fauna (though Bonaparte (1966) had tentatively suggested
that the Sinokannemeyeria fauna might be younger than this, he no longer considers
that this is likely [Bonaparte, personal communication]), the possible kannemeyeriids
Sinokannemeyeria and Parakannemeyeria are very specialized in their long snouts,
as Bonaparte (1966) has pointed out, a modification quite unlike the general trend
in dicynodont evolution.
As a result of these facts, little can be said of the history of the stahleckeriids and
286 TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
the kannemeyeriids during the Scythian or (if their differentiation had taken place
even earlier) during the late Permian. Dicynodonts with both the high, narrow
type of skull (e.g. Daptocephalus, Dinanomodon) and the low, wide type of skull
(e.g. Aulacephalodori) are found in the Upper Permian Kistecephalus Zone fauna of
South Africa. There may or may not be phyletic connections between these Permian
forms and the corresponding Triassic groups; it is, at present, impossible to decide.
Genus SANGUSAURUS nov.
DIAGNOSIS. Large dicynodont : complete skull would probably have been 35-40
cm. long. No teeth in upper or lower jaws. Premaxilla tapers anteriorly to blunt
point. Median region of intertemporal bar markedly concave in transverse section.
Low boss immediately posterior to pineal foramen. Interparietal forms posterior
end of intertemporal bar.
TYPE SPECIES. Sangusaurus edentatus sp. nov.
Sangusaurus edentatus sp. nov.
Text-figs. 21-23
HOLOTYPE. Livingstone Museum (Zambia) specimen No. LM/NH 9/1 (Field No.
15 H/4) consisting of the following damaged bones: premaxilla, right and left
maxillae, right and left quadrates; postero-dorsal corners of both squamosals ; right
prefrontal; intertemporal bar, consisting of interparietal and right and left parietals.
HORIZON AND LOCALITY. Upper fossiliferous horizon of Triassic Ntawere Forma-
tion; from locality 15 of Drysdall & Kitching (1963), about 3^ miles west of Sitwe,
in the upper Luangwa Valley, Eastern Province, Republic of Zambia.
DESCRIPTION. The material was collected as isolated fragments over an area of
about 50 square yards but, since all the bones are of commensurate size and no
individual element is duplicated, it seems very likely that all belong to a single skull.
Though much of this skull was not recovered, the fragments preserved are sufficient
to demonstrate that it belongs to a new genus of dicynodont. The generic name
Sangusaurus is derived from that of a nearby stream, the Sangu, while the specific
name edentatus refers to the lack of tusks.
The most characteristic feature of the new genus is the intertemporal bar (Text-
fig. 21). The median region of the parietals is quite deeply concave in section, as
can be seen in posterior view (Text-fig. 2ic). The sides of this trough form ridges,
parts of the lateral surfaces of which may have been originally covered by the post-
orbitals, though the bone is too badly damaged for the areas of overlap to be
distinguishable.
There is a low, median rugose boss immediately behind the pineal foramen.
The anterior ends of the parietals are damaged, but what remains of these surfaces
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
287
bears a pattern of vertical grooves similar to those found at the anterior end of the
parietal of Zambiasaurus (cf. Text-fig. le). This suggests that these surfaces are
those to which the frontals and preparietal were sutured.
The interparietal forms a wide contribution to the posterior end of the inter-
temporal bar and has a W-shaped suture with the parietals. There is an abrupt
transition between this region of the interparietal and its occipital region (Text-fig.
2ib, occ. surf, ip), since these two regions lie at a very acute angle (c. 25°) to one
another. As preserved, the occipital surface of the interparietal therefore slopes
very sharply antero-ventrally, and it seems unlikely that this is due to crushing,
since this region of the skull is extremely solid. If it is undistorted, this orientation
of the occipital surface of the interparietal implies that the whole intertemporal
pin.f.
occ. surf. ip.
FIG. 21. Sangusaurus edentatus gen. et sp. nov., intertemporal bar of type specimen, x ^.
A, dorsal view; B, lateral view; C, posterior view. Abbreviations: IP, interparietal;
occ.surf.ip., occipital surface of interparietal; P, parietal; pin.f., pineal foramen.
288
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
bar was directed postero-dorsally, so that its posterior end projected backwards
above the occiput.
The palatal surface of the premaxilla (Text-fig. 22b) bears a pair of anterior palatal
ridges. The premaxilla tapers anteriorly, so that the anterior ends of these ridges
meet the converging lateral surfaces of the bone. The grooves lateral to these
ridges therefore do not emerge along the anterior edge of the premaxilla, but along
its antero-lateral edges, and they are more shallow than the median groove which
lies between the palatal ridges. The outer surface of the anterior end of the pre-
maxilla bears a median ridge and a pair of lateral ridges.
The right maxilla (Text-fig. 3) is more complete than the left. There is no sign
of a tusk either in the caniniform processes or in either maxillary antrum. A
fairly well developed flange runs down the postero-lateral edge of the bone. The
B
FIG. 22.
saurus edentatus gen. et. sp. nov., premaxilla of holotype. x
A. lateral view; B, ventral view.
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
289
palatal surface of the maxilla is slightly recessed anteriorly where it would have been
overlapped by the premaxilla. If the maxilla is orientated so that its caniniform
process is directed vertically, the ventral edge of this recess runs antero-ventrally
at a considerable angle to the horizontal. However, the premaxilla-maxilla suture
in this region in most dicynodonts runs approximately horizontally, parallel to the
palatal surface. Such a result can be achieved only if the caniniform process is
swung forwards so as to have a very marked antero-ventral orientation.
The isolated right pref rental and a few other damaged and unidentifiable fragments
show no features of significance.
B
FIG. 23. Sangusaurus edentatus gen. etsp. nov., maxilla of holotype. x |. A, lateral view;
B, section through maxilla at a-b. Abbreviation: gr.pmx., groove marking lower limit
of area of maxilla originally covered by premaxilla.
DISCUSSION. Taxonomy and relationships of Sangusaurus. There can, first ot all,
be no doubt that Sangusaurus is generically distinct from all other dicynodonts,
since the features mentioned in the diagnosis are not found together in any previously
known genus.
Several features indicate that Sangusaurus is a kannemeyeriid rather than a
stahleckeriid. These features include the postero-dorsally directed intertemporal
bar (if the posture of this region has been interpreted correctly above) and anteriorly
taping premaxilla, the palatal ridges of which meet the lateral edges of the bone,
2Qo TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
not the anterior edge. The well developed flange down the posterior surface of the
maxilla is also commonly found in kannemeyeriids (though a small flange is present
in the stahleckeriid Zambiasaurus) .
The Kannemeyeriidae at present includes six genera: Kannemeyeria, from the
Lower Triassic Cynognathus Zone of South Africa, the Middle Triassic Manda
Formation of East Africa, and the Middle(?) Triassic Puesto Vie jo Formation of
Argentina (Bonaparte, 1966, 1967) ; Sinokannemeyeria and Parakannemeyeria, both
from the Lower Triassic Er-ma-ying Formation of Shansi, China; Barysoma, from
the Middle Triassic Santa Maria Formation of Brazil; Ischigualastia, from the Middle
or Upper Triassic Ischigualasto Formation of Argentina, and Placerias from the
Upper Triassic Chinle Formation of Arizona, U.S.A. The morphology of Ischigua-
lastia and Placerias, and the inter-relationships of all the above genera have been
discussed in an earlier paper (Cox, 1965).
Compared with the above kannemeyeriids (except Barysoma, whose skull is known
only from a partial occiput), the structure of the intertemporal bar of Sangusaurus
is closest to that of Ischigualastia. In both genera the median region of the parietals
is concave in transverse section, there is a W-shaped suture between the parietals
and the interparietal, and the posterior end of the intertemporal bar (again, if
correctly interpreted in Sangusaurus) projects posteriorly above the occiput. The
intertemporal bars of the other genera differ. That of Placerias is much wider than
that of Sangusaurus and is flatter dorsally in cross-section. That of Kannemeyeria
is very narrow, forming a sharp median crest. Those of both Sinokannemeyeria
and Parakannemeyeria are moderately wide and are concave dorsally, but are not
postero-dorsally directed, and there is therefore a more gradual transition from the
dorsal surface of the intertemporal bar on to the occiput.
Though similar to one another in general, the intertemporal bars of Sangusaurus
and Ischigualastia differ in that the interparietal is much longer in Ischigualastia
and forms the whole width of the posterior end of the intertemporal bar, and the
dorsal surface of its interparietal is convex, not concave as in Sangusaurus.
Another similarity between Sangusaurus and Ischigualastia is the lack of tusks
in both genera. All the remaining kannemeyeriids are tusked. (This region is
unknown in Barysoma, which appears to be closely related to Ischigualastia.) How-
ever, this similarity is less significant than the similarities noted above, as many
dicynodont genera have lost the tusks. If the antero-ventral orientation of the
caniniform process of Sangusaurus suggested above is correct, this is another feature
known only in this genus and in Ischigualastia.
The above comparisons suggest that Sangusaurus is more closely related to Ischi-
gualastia than to the other known Triassic kannemeyeriids. However, Sangusaurus
is so incompletely known that it would be unwise to place too much reliance upon
these resemblances, or to regard a relationship between the two genera as definitely
established.
IV. AGE OF THE NTAWERE FAUNAS
Before the significance of the faunas of the Ntawere Formation can be appreciated,
it is first necessary to establish the Triassic faunal sequence into which they must be
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 291
inserted. I shall therefore first discuss the faunas and ages of three other units:
the Cynognathus Zone of South Africa, the Molteno Beds of South Africa and the
Manda Formation of Tanganyika (Tanzania) .
1. Cynognathus Zone. This is the uppermost unit of the Beaufort Series. The
fauna includes cynognathid, diademodont and trirachodont cynodonts, the very
primitive rhynchosaurs Howesia and Mesosuchus, and the primitive pseudosuchians
Euparkeria and Erythrosuchus. The only dicynodont which has been described is
Kannemeyeria ; however, two other undescribed genera are known, one in the collec-
tion of the Bloemfontein Museum, South Africa, and the other in that of the Smith-
sonian Institution, Washington. Though the Cynognathus Zone is up to 2,000 feet
thick, this fauna is found only in the lowest 500-600 feet (Hotton & Kitching, 1963).
I have recently (Cox, 1967) suggested that the Cynognathus Zone may be best
regarded as of Upper Scythian age. This estimate was based upon the presence in
the underlying Lystrosaurus Zone (itself regarded as Lower Scythian in age) of the
capitosaurid labyrinthodont Wetlugosaurus, which had also been identified from a
bed in Greenland above a zone containing a dateable ammonite fauna. Though
Welles & Cosgriff (1965) have pointed out the unreliability of the Greenland specimens
in particular, and of this " genus " in general, Cosgriff has also provided a better
basis for the dating of the Cynognathus Zone. He has described (Cosgriff, 1965)
a new Australian temnospondyl which is associated with microfloras and marine
fossils, both of which indicate a Lower Scythian age. It is closely related to
Peltostega of the Upper Scythian Poisdonomya Beds of Spitzbergen, and to Rhytido-
steus of the Cynognathus Zone. Cosgriff (1965) also points out that comparison of
the temnospondyl amphibians of the Cynognathus Zone with those of the Lystro-
saurus Zone suggests that there was probably an appreciable time lapse between these
two faunas. Relevant here is Hotton & Kitching's (1963) comment, based on
geological interpretation, that the Cynognathus Zone may have been deposited after
a considerable interval of erosion. All these facts may be reconciled and accommo-
dated if the Lystrosaurus Zone is regarded as of Upper Permian age (as Cosgriff,
1965, suggests), and is followed by a gap in the record, equivalent to the Lower
Scythian, which is in turn followed by the Upper Scythian fauna from the lower part
of the Cynognathus Zone.
2. Molteno Beds. These beds were originally regarded as barren. Boonstra
(1947) described a cynognathid cynodont, Cynidiognathus longiceps, collected by
Stockley in what he thought were lower Molteno Beds of Basutoland (now Lesotho) .
This species is known from the Cynognathus Zone, and it is now thought possible
that this specimen may have come from nearby exposures of that Zone. More
recently, Crompton & Ellenberger (1957) described a traversodont cynodont,
Scalenodontoides, from the upper Molteno Beds ; however, Crompton (1968, personal
communication) now considers that the level from which this specimen originated
lies in the strata which are transitional between the Molteno Beds and the overlying
Red Beds, not in the typical Molten Beds. As a result, once again, vertebrate
fossils are unknown from the Molteno Beds. These beds, up to 1,900 feet thick,
may therefore cover all or part of the time between the underlying Upper Scythian
292 TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
Cynognathus Zone and the overlying Red Beds. These latter contain a varied
dinosaur fauna, including melanorosaurs, thecodontosaurs and sauropod-like foot-
prints, and are probably of Upper Triassic age (Charig, Attridge & Crompton, 1965).
No more accurate assessment can at present be made than that the Molteno Beds
therefore represent part or all of the Middle Triassic (Ladinian and Anisian).
3. Manda Formation. This fauna lacks the more primitive cynognathid and
diademodont cynodonts found in the Cynognathus Zone, but instead includes
traversodont cynodonts. Both the rhynchosaur and the diverse pseudosuchians are
far more advanced and common than are those of the Cynognathus Zone. The
dicynodonts include a greater variety of forms than the Cynognathus Zone, but the
genus Kannemeyeria is still present. The Manda fauna has provisionally been
regarded as of Anisian (possibly Upper Anisian) age (Cox, 1965, 1967).
4. N taw ere Formation. As already mentioned, this formation contains two
fossiliferous horizons, which will be discussed separately. That from the lower part
of the formation contains two undescribed new tusked dicynodonts (in the collection
of the Bernard Price Institute for Palaeontological Research, Johannesburg), the
cynodont Diademodon rhodesiensis , and fragments of large labyrinthodonts. The age
of this fauna must at present depend upon the characters of the cynodont. Brink
(1963 : 79) states that this is " very typically a Diademodon and it is very difficult
to find a particular feature about the skull on the strength of which it can emphatically
be differentiated from a known species like D. polyphagus " (of the Cynognathus
Zone). Brink notes, however, that the post-dentary bones are more reduced,
and the coronoid process of the dentary better developed, than in the Cynognathus
Zone species. These somewhat advanced features may indicate that this Ntawere
fauna is slightly younger than that of the Cynognathus Zone, and it might conveni-
ently be regarded as equivalent to the upper, non-fossiliferous, portion of the
Cynognathus Zone.
The upper fossiliferous horizon extends over the topmost beds of the Ntawere
Formation and the lower part of the Red Marl. Its fauna includes the stahleckeriid
dicynodont Zambiasaurus, the kannemeyeriid dicynodont Sangusaurus, two
traversodont cynodonts (Luangwa drysdallensis Brink 1963 and another specimen
which will be described by Crompton), fragments of pseudosuchians, and the remains
of the largest known labyrinthodont amphibian, approximately twice the size of
Paracydotosaurus (which will be described by Panchen).
The absence of diademodonts (as far as is known), the presence of traversodonts
and its stratigraphical position above the lower fossiliferous horizon (itself of possible
Cynognathus Zone affinities) all indicate that this upper Ntawere fauna is younger
than the Cynognathus Zone. The above characters of its cynodont fauna similarly
indicate a closer relationship to the Manda fauna. The only remaining problem is
whether this Ntawere Fauna should be regarded as older than, or the same age as,
the Manda Formation. The only fact relevant to this decision is the absence from
the Ntawere fauna of the rhynchosaurs which form such a large element in the Manda
fauna. Though, as has been pointed out previously (Cox, 1967), this group is notably
unreliable as a faunal indicator, it must be accepted here in the absence of any more
TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA 293
reliable evidence. The upper fossiliferous horizon of the Ntawere Formation is
therefore provisionally to be regarded as slightly older than the Manda Formation.
If this is so, the suggestion of Dixey (1936) and of Drysdall & Kitching (1964) that
the Escarpment Grit (which underlies the Ntawere Formation) may be equivalent
to the Kingori Sandstone (which underlies the Manda Formation) must be incorrect
unless the Kingori Sandstone is equivalent to both the Ntawere Formation and the
Escarpment Grit. It is, perhaps, more likely that the Kingori Sandstone represents
a later phase in the sedimentary cycle which began with the Escarpment Grit.
TABLE i
South Africa Zambia Tanzania
Rhaetian Cave Sandstones
Norian Red Beds
Carnian "1
Ladinian > Molteno Beds
U. Anisian J Manda Formation
L. Anisian Unfossiliferous Ntawere Formation Kingori Sandstone
U. Cynognathus Zone
U. Scythian Cynognathus Zone Escarpment Grit
fauna
L. Scythian
U. Permian Lystrosaurus Zone
Kistecephalus Zone Madumabisa Kawinga Formation
Mudstone
V. ACKNOWLEDGMENTS
Most of the expenses of the 1963 expedition, during which these specimens were
collected, were met by grants from the Department of Scientific and Industrial
Research and the Royal Society ; grants were also kindly made by the Percy Sladen
Memorial Fund, the Godman Exploration Fund and Shell Research Ltd. All the
members of the Expedition are grateful to these bodies for their support, which made
the Expedition possible.
The drawings in this paper are by Miss J. Joffe and Mr. P. Hutchinson, who were in
turn employed as my research assistant. The salary of this post was met from a
special grant of the Natural Environment Research Council, for whose support I am
greatly indebted.
VI. REFERENCES
ATTRIDGE, J., BALL, H. W., CHARIG, A. J. & Cox, C. B. 1963. The British Museum (Natural
History) — University of London Joint Palaeontological Expedition to Northern Rhodesia
and Tanganyika, 1963. Nature, Lond. 201 : 445-449, 4 figs.
BONAPARTE, J. F. 1966. Una nueva " fauna " Triasica de Argentina (Therapsida: Cyno-
dontia Dicynodontia) . Consideraciones filogeneticas y paleobiogeograficas. Ameghiniana,
Buenos Aires, 4 : 243-296, 29 figs., 2 pis.
— 1967. New vertebrate evidence for a southern transatlantic connexion during the Lower
or Middle Triassic. Palaeontology, London, 10 : 554-563, 7 figs.
294 TWO NEW TRIASSIC DICYNODONTS FROM ZAMBIA
BOONSTRA, L. D. 1947. Notes on some Stormberg fossil bones from Basutoland. In G. M.
Stockley's Report on the geology of Basutolani, pp. 94-95, i pi. Maseru.
BRINK, A. S. 1963. Two cynodonts from the Ntawere Formation in the Luangwa Valley of
Northern Rhodesia. Palaeont. afr., Johannesburg, 8 : 77-96, figs. 12-15.
CAMP, C. L. 1956. Triassic dicynodont reptiles. Part II. Triassic dicynodonts compared.
Mem. Univ. Calif., Berkeley, 13 : 305-341, figs. 42-62.
CHARIG, A. J., ATTRIDGE, J. & CROMPTON, A. W. 1965. On the origin of the sauropods
and the classification of the Saurischia. Proc. Linn. Soc. Lond. 176 : 197-221.
COSGRIFF, J. W. 1965. A new genus of Temnospondyli from the Triassic of Western Australia.
/. Proc. R. Soc. West. Aust., Perth, 48 : 65-90, 13 figs.
Cox, C. B. 1965. New Triassic dicynodonts from South America, their origins and relation-
ships. Phil. Trans. R. Soc., London, 248B : 457-516, 30 figs.
— 1967. Changes in terrestrial vertebrate faunas during the Mesozoic. In: Harland, W. B.
et al. eds. The Fossil Record, London (Geological Society), pp. 77-89, i fig.
— 1968. The Chanares (Argentina) Triassic reptile fauna. IV. The dicynodont fauna.
Breviora, Cambridge, Mass. No. 205: 1-27, 12 figs.
CROMPTON, A. W. & ELLENBERGER, F. 1957. On a new cynodont from the Molteno Beds and
the origin of the tritylodontids. Ann. S. Afr. Mus., Cape Town, 44 : 1-14, 5 figs., i pi.
CROMPTON, A. W. & HOTTON, N. 1967. Functional morphology of the masticatory apparatus
of two dicynodonts (Reptilia, Therapsida). Postilla, New Haven, No. 109 : 1-51, 7 figs.
DIXEY, F. 1936. The Karroo of the upper Luangwa valley of north-east Rhodesia. Rep.
geol. Surv. Dep. Nyasald., Livingstone, 24 :
— 1937. The geology of part of the upper Luangwa valley, north-eastern Rhodesia. Q. Jl
geol. Soc. Lond. 93 : 52-74, i fig., pi. iv.
DRYSDALL, A. R. & XITCHING, J. W. 1963. A re-examination of the Karroo succession and
fossil localities of part of the upper Luangwa Valley. Mem. geol. Surv. Dep. N. Rhod.,
Lusaka, 1 : 1-62, 4 figs. 7 pis.
HOTTON, N. & KITCHING, J. W. 1963. Speculations on Upper Beaufort deposition. S. Afr.
J. Sci., Cape Town, 59 : 254-258.
HUENE, F. VON 1935-42. Die fossilen Reptilien des sudamerikanischen Gondwanalandes an der
Zeitenwende. 332 pp., 66 figs., 38 pis. Tubingen.
WELLES, S. P. & COSGRIFF, J. W. 1965. A revision of the labyrinthodont family Capito-
sauridae. Univ. Calif. Publs. Bull. Dep. Geol., Berkeley, 54 : 1-148, 48 figs., i pi.
PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING
LOWER CAMBRIAN L2S
ARCHAEOCYATHA FROM THE
AJAX MINE,
BELTANA, SOUTH AUSTRALIA
FRANCOISE DEBRENNE
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 7
LONDON: 1969
LOWER CAMBRIAN ARCHAEOCYATHA
FROM THE AJAX MINE,
BELTANA, SOUTH AUSTRALIA
BY
FRANCOISE DEBRENNE
Museum National d'Histoire Naturelle, Paris
Pp. 295-376; 18 Plates, 15 Text-figures.
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 7
LONDON: 1969
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 17, No. 7 of the Geological
(Palaeontological) series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation :
Bull. Br. Mus. nat. Hist. (Geol.).
Trustees of the British Museum (Natural History) 1969
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 15 April, 1969 Price £4 155.
LOWER CAMBRIAN ARCHAEOCYATHA
FROM THE AJAX MINE,
BELTANA, SOUTH AUSTRALIA
By FRANCOISE DEBRENNE
CONTENTS
Page
I. INTRODUCTION ......... 299
II. SYSTEMATIC DESCRIPTIONS ....... 302
Phylum ARCHAEOCYATHA . . . . . .302
Class REGULARIA Vologdin, 1937 • • • • • 3°2
Order MONOCYATHIDA Okulitch, 1935 .... 302
Family MONOCYATHIDAE R. & W. R. Bedford, 1934 • 3°2
Genus MONOCYATHUS R. & W. R. Bedford, 1934 302
M. porosus R. & W. R. Bedford . . . 303
M. mellifer R. & W. R. Bedford ... 303
Genus TUMULIOLYNTHUS Zhumvlevz, 1963 . 304
T. irregularis (R. & W. R. Bedford) . . 304
Order AJACICYATHIDA R. & J. Bedford, 1939 . . . 305
Family DOKIDOCYATHIDAE R. & W. R. Bedford, 1936 . 305
Genus ALPHACYATHUS R. & J. Bedford, 1939 . 305
A. cf. annularis (R. & W. R. Bedford) . . 305
Family ACANTHINOCYATHIDAE R. & W. R. Bedford, 1936 306
Genus ACANTHINOCYATHUS R. & W. R. Bedford, 1936 306
A. apertus R. & J. Bedford .... 307
Family AJACICYATHIDAE R. & J. Bedford, 1939 . . 308
Genus ARCHAEOCYATHELLUS Ford, 1873 . 308
Subgenus STAPICYATHUS Debrenne, 1964 . 309
A. (S.) stapipom (Taylor) . . . 309
Genus LOCULICYATHUS Vologdin, 1931 . . 310
Subgenus LOG ULICYATHELLUS nov. . . 310
L. (L.) floreus (R. & W. R. Bedford) . 310
Family ROBUSTOCYATHIDAE Debrenne, 1964 . . . 311
Genus ROBUSTOCYATHUS Zhuravleva, 1960 . 311
R. magnipora (R. & W. R. Bedford) . . 311
R. subacutus (R. & W. R. Bedford) . . 312
Genus ZONACYATHUS R. & J. Bedford, 1937 . 314
Z. retevallum (R. & W. R. Bedford) . . 315
Z. retezona (Taylor) . . . . . 316
Family BRONCHOCYATHIDAE R. & J. Bedford, 1936 . 317
Genus CYATHOCRICUS nov 318
C. tracheodentatus (R. & W. R. Bedford) . 319
C. dentatus (Taylor) . . . . .321
Family ETHMOCYATHIDAE nov. ..... 322
Genus ETHMOCYATHUS R. & W. R. Bedford, 1934 323
E. lineatus R. & W. R. Bedford ... 323
GEOL. 17, 7 24
298 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
Family ERISMACOSCINIDAE Debrenne, 1964 . . . 325
Genus ERISMA COSCIN US Debrenne, 1958 . . 325
E. rugosus (R. & W. R. Bedford) . . . . . 325
E. equivallum (Taylor) . . . . 326
E. quadratus (R. & W. R. Bedford) . . 328
E. textilis (R. & W. R. Bedford) ... 329
E. cellularis (R. & W. R. Bedford) . . 330
E. peter si (R. & W. R. Bedford) . . . 331
E. retifer (R. & W. R. Bedford) ... 332
Family POLYCOSCINIDAE Debrenne, 1964 . . . 334
Genus TOMOCYATHUS Rozanov, 1960 . . 334
Subgenus ERUGATOCYATHUS nov. . . 334
T. (E.) papillatus (R. & W. R. Bedford) 334
Family SALAIROCYATHIDAE Zhuravleva, 1955 . . 336
Genus SALAIROCYATHUS Vologdin, 1940 . . 336
Subgenus S. (SALAIROCYATHUS) Vologdin, 1940 336
? S. (S.) annulatus (R. & W. R. Bedford) 337
Family ALATAUCYATHIDAE Zhuravleva, 1955 . . 338
Subfamily TUMULOCOSCININAE Zhuravleva, 1960 . 338
Genus ETHMOCOSCINUS Simon, 1939 . . 338
E. papillipom (R. & W. R. Bedford) . . 339
Subfamily ALATAUCYATHINAE Zhuravleva, 1960 . . 340
Genus ALATAUCYATHUS Zhuravleva, 1955 . 340
Subgenus A NAPTYCTOCYATHUS nov. . . 340
A . (A.) cribripora R. & W. R. Bedford . 340
A. (A.) flabellus sp. nov. . . . 341
Class IRREGULARIA Vologdin, 1937 ..... 342
Order ARCHAEOCYATHIDA Okulitch, 1935 . . . . 342
Family DICTYOCYATHIDAE Taylor, 1910 . . . 342
Genus PIN A COC YA TH US R. & W. R. Bedford, 1 934 342
P. spicularis R. & W. R. Bedford. . . 343
Family FLINDERSICYATHIDAE R. & J. Bedford, 1939 . 344
Genus FLINDERSICYATHUS R. & J. Bedford, 1937 344
Subgenus FLINDERSICYATHUS R. & J.
Bedford, 1937 . • • • • 34^
F. (F.) graphicus (R. & W. R. Bedford) . 346
F. (F.) irregularis (Taylor) . . . 348
F. (F.) major (R. & W. R. Bedford) . 349
F. (F.) speciosus (R. & W. R. Bedford) . 350
Subgenus PYCNOIDOCYATHUS Taylor, 1910 . 350
F. (P.) synapticulosus (Taylor) . . 351
F. (P.) simplex (Taylor) . . . 352
F. (P.) maximipora (R. & W. R. Bedford) 352
F. (P.) vicinisepta (R. & W. R. Bedford) . 353
Family METACYATHIDAE R. & W. R. Bedford, 1934 • 354
Genus METALDETES Taylor, 1910 ... 355
M. dissepimentalis (Taylor) .... 358
M. irregularis (R. & W. R. Bedford) . . 359
M. taylori (R. & W. R. Bedford) ... 360
Genus METAFUNGIA R. & W. R. Bedford, 1934 . 361
M. reticulata R. & W. R. Bedford . . 362
Family METACOSCINIDAE R. & W. R. Bedford, 1936 . 363
Genus METACOSCINUS R. & W. R. Bedford, 1934 3^3
M. reteseptatus R. & W. R. Bedford . . 364
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 299
? Family METACOSCINIDAE ...... 367
Genus PYCNOIDOCOSCINUS R. & W. R. Bedford,
1936. . .... 367
P. pycnoideum R. & W. R. Bedford . . 367
III. STRATIGRAPHICAL CONCLUSIONS . ...... 368
IV. SUMMARY .......... 371
V. ACKNOWLEDGMENTS ........ 372
VI. REFERENCES .......... 373
APPENDIX GLOSSARY ....... 376
SYNOPSIS
Specimens from the Ajax Mine, first described by R. & W. R. Bedford (1934, J936) and now in
the British Museum (Natural History), have been re-examined and are revised. Fifteen families,
21 genera, 4 subgenera and 41 species are discussed and arranged according to current classifica-
tion. One family (Ethmocyathidae), one genus (Cyathocricus) , three subgenera (Loculicyathellus,
Erugatocyathus, Anaptyctocyathus) and one species (Anaptyctocyathus flabellus) are considered
new. The two genera Metacyathus and Bedfordcyathus are shown to be synonyms of Metaldetes.
The excellent preservation of this silicified material has enabled the internal structures of some
species to be determined accurately and resulted in the discovery of new wall and intervallum
types.
Stratigraphical correlation shows that there are clear affinities between the Ajax, Kameshki
and Sanashtygkol faunas. The presence of both advanced and simple forms in the Ajax fauna
indicates that it is probably of Upper Kameshki-Lower Sanashtygkol age, i.e. the middle of the
lower division of the Lower Cambrian.
Ce travail a pour but de reviser les Archeocyathes qui firent 1'objet du premier m6moire de
R. et W. R. Bedford (1934) et qui sont actuellement conserv6s dans les collections du B.M.(N.H.).
15 families, 21 genres, 4 sous-genres et 41 especes sont decrits et classes selon les criteres actuels
de classification. Une nouvelle famille (Ethmocyathidae), un nouveau genre (Cyathocricus),
trois sous-genres (Loculicyathellus, Erugatocyathus, Anaptyctocyathus} et une nouvelle espece
ont ete etablies. Bedfordcyathus et Metacyathus tombent en synomymie avec Metaldetes. La
fossilisation exceptionnelle du materiel silicifi6 a permis la definition de nouvelles structures
murales et intervallaires.
Les faunes d'Ajax ont des affinites avec les faunes des horizons de Kameshki et Sanashtygkol,
c'est a dire qu'elles datent du milieu de la partie inferieure du Cambrien inferieur.
I. INTRODUCTION
THE present work is a revision of material in the British Museum (Natural History)
collection, collected and originally described by R. & W. R. Bedford in 1934. The
specimens come from a Lower Cambrian exposure in a limestone hill behind the Ajax
copper mine, ten miles north-east of Beltana, in the Flinders Range, South Australia.
T. G. Taylor was the first to visit this locality in February, 1906 and his subsequent
monograph on Australian Archaeocyatha (1910) was the first work to give detailed
descriptions of such fossils.
In a series of papers from 1934-1939, the Bedfords, R., W.R. & J., described
material collected in South Australia. The specimens mentioned in their papers now
belong to the following museums, according to correspondence in the British Museum
(Natural History) from Dr. Dorothy Hill: Memoir I — B.M. (N.H.) ; Memoir 2 — South
Australian Museum ; Memoirs 3, 4 and 6 — Princeton University, U.S.A. The remain-
300 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
Unmefarrtorpriosec/ racers,
sands/ones, sh»/rs.
CAMBRIAN OUTCROPS
OF
SOUTH AUSTRALIA
»vi»h sliqht modifications from the
Geological Map of South Australia
D«p». of Min«s, Adelaide. 1953
30°
FIG. i. The Cambrian outcrops of South Australia. After B. Daily, 1956.
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 301
der of the Bedford collection was acquired by Prof. D. A. Brown of the Geology Dept,
Australian National Museum, Canberra.
The silicified Australian material enabled both Taylor and the Bedfords to discover
details of the finer internal structure by etching their specimens. Some credit is due
to the Bedfords for the classification of the Archaeocyatha and their recognition of the
systematic importance of ontogeny in such a classification.
FIG. 2. Section from a generalized geological map of the Flinders Ranges, produced by the
Geol. Survey of South Australia and published in Glaessner, M. F. & Parkin, L. W. 1958.
The study of Archaeocyatha has figured prominently in assigning a Cambrian age to
these rocks. Daily (1956) recognized 5 faunal assemblages in the Ajax limestones, but
considered that the material collected by both Taylor and the Bedfords could only
come from his faunal assemblage No. I, which is given a Lower Cambrian age. Yet
Walter (1967) states that the Ajax fauna cannot be placed in Daily's scheme of faunal
units.
Recently Dorothy Hill (1965), in her work on Antarctic Archaeocyatha, revised the
phylum and provided good illustrations of type material in Adelaide University, the
South Australian Museum, Princeton University and the British Museum (Nat. Hist.).
At her suggestion, I undertook the study of the Bedford material in the latter collec-
tion, with the intention of providing further description, re-classification and, above
all, better figures of the specimens. It has often been difficult in the past to assess the
characters of genera and species mentioned by the Bedfords, as their figures are either
composite, or cannot be located on the actual specimen. The genus Syringocnema is
not dealt with in this paper, although well represented in that part of the Bedford
collection in the B.M. (N.H.), because of its thorough description by both Taylor
(1910 : 153) and Gordon (1920 : 699).
302 LOWER CAMBRIAN ARCH AEOCY ATM A FROM AUSTRALIA
" Archaeocyatha are known from all the continents except South America and are
characteristic of the calcareous facies of the Lower Cambrian " (Hill 1965 : 30). This
widespread distribution makes them particularly useful for stratigraphical correla-
tion; yet, as Walter (1967) points out, their use is limited for, being benthonic,
dispersal is restricted. " Recognition of the extinct Archaeocyatha as a separate
phylum near the Porifera and Coelenterata, is now general " (Hill 1965 : 45). In the
past, their systematic position has been the subject of considerable debate, with
opinion ranging from calcareous algae to sponges. However, they are regarded by
Hill (1964 : 253) " as single multicellular organisms, with organization higher than
that of the Protozoa, but with less differentiation than the Porifera ".
A short glossary of the more important descriptive, morphological terms is included
as an Appendix.
II. SYSTEMATIC DESCRIPTIONS
Phylum ARCHAEOCYATHA Vologdin, 1937
Class REGULARIA Vologdin, 1937
Order MONOCYATHIDA Okulitch, 1935
Family MONOCYATHIDAE R. & W. R. Bedford, 1934
DIAGNOSIS. Single-walled cup. Pore system simple to slightly complex.
COMPOSITION OF THE FAMILY. Monocyathus Bedford, R. & W.R. 1934, Rhabdolyn-
thus Zhuravleva 1960, Tumuliolynthus Zhuravleva 1963, ? Tunkia Bedford, R. & J.
1936.
Genus MONOCYATHUS R. & W. R. Bedford, 1934
1899 Rhabdocyathus von Toll (non Brooks 1893) : 45, pi. 8, figs. 2c, 6 and 7, t.-figs. 4-7.
1910 Archaeolynthus Taylor : 157, pi. 5.
1934 Monocyathus R. & W. R. Bedford : 2, pi. i, fig. i.
1936 Monocyathus R. & W. R. Bedford : 12, pi. 10, fig. 46.
1937 Rhabdocnema Okulitch : 251
1939 Monocyathus R. & W. R. Bedford; R. & J Bedford : 69, pi. 42, fig. 161.
1939 Archaeolynthus Taylor; Simon : 21.
1949 Archaeolynthus Taylor; Zhuravleva : 549.
TYPE SPECIES. Monocyathus porosus R. & W. R. Bedford, 1934, selected by R. &
W. R. Bedford, 1936.
DISCUSSION. Okulitch (1950), Debrenne (1964) and Hill (1965) consider Archaeo-
lynthus Taylor 1910 an invalid name, as the type specimen was not designated by the
author and the reference material was destroyed by serial sectioning.
DIAGNOSIS. Small conical cups with a simple porous single wall and the vertical
rows of pores in a quincunx pattern.
COMPOSITION OF THE GENUS. Monocyathus absolutus (Vologdin 1940), M. bilateralis
(Vologdin 1962), M. contractus Hill 1965, M. copulatus (Vologdin 1940), M. kuzneskii
(Vologdin 1931), M. lebedevae (Vologdin 1937), M. macrospinosus (Zhuravleva 1963),
M. mellifer R. & W. R. Bedford, 1936, M. nalivkini (Vologdin 1939), M. operculatus
LOWER CAMBRIAN ARCH AEOC Y ATM A FROM AUSTRALIA 303
Maslov 1960, M. partibus (Vologdin 1963), M. polaris (Vologdin 1937), M. porosus
R. & W. R. Bedford, 1934, M. robustus R. & W. R. Bedford, 1936, M. sibiricus (von
Toll 1899), M. simplex (Vologdin 1940), M. sparsipom R. & W. R. Bedford, 1936, M.
spinosus R. & W. R. Bedford, 1936, M. tenuimurus (Vologdin 1940), M. tolli (Krasno-
peeva 1937), M. unimurus (Vologdin 1940).
Monocyathus porosus R. & W. R. Bedford
(PL i, ng. 3)
1934 Monocyathus porosus R. & W. R. Bedford : 2, pi. i, fig. i.
1939 Monocyathus porosus R. & J. Bedford : 69, fig. 161.
1963 Archaeolynthus porosus (Bedford & Bedford) ; Zhuravleva : 88-89, fig. 39.
1965 Monocyathus porosus R. & W. R. Bedford; Hill : 52, pi. 2, fig. i.
LECTOTYPE. B.M. (N.H.) S 4140 selected by Hill (1965).
OTHER MATERIAL. Syntypes B.M. (N.H.) S 7630-31, S 4783-8.
DESCRIPTION. Conical cup, slightly waved, with a single wall of constant thickness
and pores in quincunx. The upper edge, in the material studied, is not turned back
into the central cavity as an incipient pelta. The pores are proportionately smaller
and more numerous towards the upper part of the cup. The bigger specimens have
smaller and more regular pores.
DIMENSIONS
Lectotype S 7630-7631
(mm.) (mm.)
Height (pars) 15 31
Diameter 6 9-5
Wall:
Diameter of the pores o • 27-0 -20 o • 10
Distance between the vertical rows 0-33-0-27 °'33
Distance between the horizontal rows o • 54-0 -67 o • 35
DISCUSSION. This species is very similar to the Siberian species M. nalivkini
(Vologdin). Zhuravleva (1963 : 80) distinguishes the two, mainly on the supposed
presence of a pelta and inner rugosity in porosus, which was described and figured by
R & W. R. Bedford (1936). However, these peculiar structures are neither seen in
the lectotype, nor on the other specimens in the British Museum collection. It is
therefore possible that the two species are synonymous.
Monocyathus mellifer R. & W. R. Bedford
(PI. i, fig. 4)
1936 Monocyathus mellifer R. & W. R. Bedford : 12, pi. 10, fig. 49.
MATERIAL. B.M. (N.H.) S 4821. The whereabouts of the holotype is not known.
REMARKS. The specimen is a cylindrical fragment of a single-walled cup. The
wall is like a honeycomb, formed by short hexagonal pipes with a hexagonal section
on the outer side and a circular section on the inside.
304 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
DIMENSIONS
(mm.)
Height (pars) 16
Diameter 3 • 72
Wall:
Inner diameter of pores o • 40
Pipe wall thickness 0-27
Wall thickness 0-67
DISCUSSION. This fragmentary piece is similar to the type-species briefly described
by R. & W. R. Bedford 1936.
Genus TUMULIOLYNTHUS Zhuravleva, 1963
1932 Rhabdocyathus von Toll : Vologdin (pars) : 65.
1963 Tumuliolynthus Zhuravleva : 101.
TYPE SPECIES. Rhabdocyathus tubexternus Vologdin 1932, by original designation
Zhuravleva (1963 : 101).
DIAGNOSIS. One-walled Archaeocyatha, with the pores protected externally by
simple tumuli.
COMPOSITION OF THE GENUS. (After Zhuravleva 1963) : T. irregularis (R. & W. R.
Bedford, 1934), T. karakolensis Zhuravleva 1963, T. musatovi (Zhuravleva 1961),
T. tubexternus (Vologdin 1932), T. vologdini (Yakovlev 1956).
Tumuliolynthus irregularis (R. & W. R. Bedford)
(PI. i, fig. i)
1934 Monocyathus irregularis R. & W. R. Bedford : 2, pi. i, fig. 2.
1939 Monocyathus irregularis R. & W. R. Bedford; R. & J. Bedford : 68, fig. 160.
1963 Tumuliolynthus irregularis (R. & W. R. Bedford) Zhuravleva : no, fig. 58.
HOLOTYPE. B.M. (N.H.) S 4141.
OTHER MATERIAL. B.M. (N.H.) S 7643-6, S 4771, S 4774-7, S 4764-5.
REMARKS. The holotype is a small cylindrical fragment with several large aper-
tures, each of which is probably the basal trace of a pore. The scattered tumuli
occur on the lower parts and have a large opening at the top. The poor preservation
and small size of this specimen prevent a more detailed description. The other
specimens are more complete, generally larger, and have a thicker wall with irregular
apertures, each with a tumulus perforated at the top.
DIMENSIONS
S 4141 S 4774-4777 S 4817-4820
(mm.) (mm.) (mm.)
Cup:
Height 6-1 18
Diameter 1-69 0-33 0-27
Wall:
Diameter of pores o • 60-0 -23 0-20 0-27
Average distance between pores . . 0-67
Thickness o-io 0-33 0-27
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 305
DISCUSSION. R. & W. R. Bedford, (1934) and R. & J. Bedford, (1939) noted the
presence of " papillae " and only placed this species in the genus Monocyathus
provisionally. Zhuravleva (1963) uses the term " tumulus " when referring to the
papillae and after examining the material, I consider her change of terminology
justified.
Tumuliolynthus irregularis differs from other species of the genus by the irregular
size and pattern of its pores.
Order AJACICYATHIDA R. & J. Bedford, 1939
Family DOKIDOCYATHIDAE R. & W. R. Bedford, 1936
DIAGNOSIS. Two-walled cup, walls simply porous, connected by radial horizontal
rods.
COMPOSITION OF THE FAMILY, Dokidocyathus Taylor 1910, Alphacyathus R. & J.
Bedford, 1939.
Genus ALPHACYATHUS R. & J. Bedford
!Q39 Alphacyathus R. & J. Bedford : 72.
TYPE SPECIES. Dictyocyathus annularis R. & W. R. Bedford, 1936, by original
designation R. & J. Bedford, (1939 : 72, fig. 55).
DIAGNOSIS. Cup with two simply porous walls. In the intervallum, radial
cylindrical bars are arranged in regularly spaced horizontal planes and are connected
by synapticulae that are opposite from loculus to loculus, forming a continuous
ring in the centre of the intervallum.
COMPOSITION OF THE GENUS. A single species Alphacyathus annularis (R. & W. R.
Bedford, 1936).
Alphacyathus cf. annularis (R. & W. R. Bedford)
(PI. I, fig. 2)
cf. 1936 Dictyocyathus annularis R. & W. R. Bedford : 13, pi. u, fig. 55.
1939 Alphacyathus annularis R. & W. R. Bedford) R. & J. Bedford : 72.
HOLOTYPE. P 942 in the South Australian Museum, Adelaide according to Hill
(1965 : 55)-
OTHER MATERIAL. B.M. (N.H.) S 4822, S 4766.
DESCRIPTION. Small cylindrical cups. Outer wall is pierced by circular pores in
quincunx, the skeletal tissue between the pores (" linteaux ") is of constant thickness
and equal in width to the pore diameter. Inner wall is built on the same pattern with
regular pores and considerable skeletal tissue. One inner wall pore at each inter-
radial space and at each horizontal level.
306 LOWER CAMBRIAN ARCH AEOCY ATM A FROM AUSTRALIA
DIMENSIONS
S 4822 S 4766
(mm.) (mm.)
Cup:
Height ....... 5 unknown
Diameter ....... 2 2 • 45
Intervallum . . . . . . 0-70 0-50
Central cavity . . . . . . 0-56 0-90
Outer wall:
No. of pores between 2 bars . . . 2-3 2-3
Diameter . . . . . . . 0-07 0-07
Vertical partitions . . . . . 0-07 0-07
Horizontal partitions . . . . . 0-07 0-07
Thickness . . . . . . . 0-07 0-07
Inner wall :
No. of pores between 2 bars i i
Diameter . . . . . . . 0-15 0-13
Vertical partitions . . . . . 0-07 0-15
Horizontal partitions . . . . . 0-07 0-15
Thickness ....... 0-07 0-15
Rods:
Interradial space . . . . . . . 0-2
Vertical space . . . . . .0-2
Diameter . . . . . . . 0-15 0-15
DISCUSSION. Neither specimen provides a sufficiently complete transverse section
for showing the synapticulae and they cannot be assigned with certainty to annular is.
Family ACANTHINOCYATHIDAE R. & W. R. Bedford, 1936
DIAGNOSIS. Two-walled cups with intervallar horizontal rods. Inner wall simple,
but outer wall has protected pores, the lower skeletal part of each pore bearing a long
spinous process, projecting upwards and outwards from the wall.
COMPOSITION OF THE FAMILY. Acanthinocyathus R. & W. R. Bedford, 1934.
Genus ACANTHINOCYATHUS R. & W. R. Bedford, 1936
1934 Acanthocyathus R. & W. R. Bedford (non Edwards & Haime 1848, which is a hexacoral) : 4.
1936 Acanthinocyathus R. & W. R. Bedford : n.
TYPE SPECIES. Acanthocyathus apertus R. & W. R. Bedford (1934 : 4, fig. 20), by
monotypy.
DIAGNOSIS. Two-walled cups with radial, horizontal, or upwardly oblique rods.
The outer wall pores are partly obscured by scales. The inner wall consists of a net of
polygonal to circular pores.
COMPOSITION OF THE GENUS. A. apertus (R. & W. R. Bedford), ? A. transiens
R. & J. Bedford 1939.
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA 307
Acanthinocyathus apertus R. & W. R. Bedford
(PI. 2, Text-fig. 3)
1934 Acanthocyathus apertus R. & W. R. Bedford : 4, fig. 20.
1936 Acanthinocyathus apertus (R. & W. R. Bedford); R. & W. R. Bedford : n, fig. 45.
LECTOTYPE. B.M. (N.H.) S 4166 chosen here.
OTHER MATERIAL. Para-lectotypes B.M. (N.H.) S 4167-8.
DESCRIPTION. Cylindrical rods cross the intervallum and are orientated horizont-
ally, or sometimes obliquely upwards and outwards from the inner to the outer wall.
FIG. 3. Acanthinocyathus apertus R. & J. Bedford
The inner wall is well shown in S 4168 ; it consists of a large pored net, with a mesh of
irregular size and shape formed by skeletal threads of constant thickness. The outer
wall is the most characteristic feature of this species ; the large pores are in quincunx
and of various shapes ; the skeletal tissue occupies a smaller area than the pores and
carries long spines that are directed upwards and outwards. Unfortunately, these
long protective spines are often broken, but when complete they may reach the
centre of the overlying pore, that is to say, since the rows alternate, the centre of the
pore two rows above. The base of the spine is level with the lower third of the pore.
308 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
DIMENSIONS
Holotype
S 4166 S 4168 S 4167
(mm.) (mm.) (mm.)
Cup:
Height (pars) 34 32 25
Upper diameter 1 7 • 5 13 1 1
Lower diameter 15 12-5 8-5
Upper intervallum 3-5 3-4 3
Lower inter vallum 3-4 3-4 2-37
Outer wall:
Diameter of pores i -7-2 . . i -oi-i -35
scales 4 • 75-6 . . 3-4
Vertical partitions i • 35 . . 0-67
Horizontal partitions 1-35 . . 1-35
Thickness 0-67 . . 0-6
Inner wall:
Diameter of pores 0-75 0-2 7-2 -03 0-6-1-15
Partitions + 0-61 0-4
Thickness 0-40 0-47 0-4
Rods 0-4
DISCUSSION. R. & W. R. Bedford, thought the inner wall was a scaffolding of tri-
radiate spicules, the side rays surrounding the pores and the median ray being pro-
longed into a spinous process directed upwards and outwards. They considered that
the spicular elements of Acanthinocyathus could indicate a phylogenetic link between
Sponges and Archaeocyatha. This suggestion is not borne out by observation. The
wall of Acanthinocyathus is a porous sheet with the elongation of its horizontal skeletal
parts into scales of an unusual size.
Family AJACICYATHIDAE R. & J. Bedford, 1939
Genus ARCHAEOYCATHELLUS Ford, 1873
Subgenus STAPICYATHUS Debrenne, 1964
1873 Archaeocyathellus Ford (Pars).
1964 Archaeocyathellus (Stapicyathus) Debrenne : 127.
TYPE SPECIES. Archaeocyathus stapipora Taylor (1910 : 118), by original designa-
tion of Debrenne (1964 : 127).
DIAGNOSIS. Conical cups; outer and inner walls with simple pores as in true
Ajacicyathidae. Intervallum crossed by imperf orate radial septa. The neighbour-
ing loculi only open into one another and into the central cavity, by the stirrup-pores
of the inner wall.
REMARKS. Debrenne (1964 : 127) considered this form a non-corrugated subgenus
of Archaeocyathellus Ford.
COMPOSITION OF THE SUBGENUS. A. (S.) stapipora (Taylor 1910) and ? A. (S.)
yukonensis (Okulitch 1957).
LOWER CAMBRIAN ARCH AEOC Y ATM A FROM AUSTRALIA
309
Archaeocyathellus (Stapicyathus) stapipora (Taylor)
(PI- i, %. 5)
1910 Archaeocyathus stapipora Taylor : 118, pi. 7, figs. 37 and 38, pi. 3, fig. 10, t-fig. 14.
1939 Archaeocyathus stapipora Taylor; R. & J. Bedford : 75.
IQ39 Archaeocyathus (Protocyathus) stapipora (Taylor) Simon : 54.
1964 Archaeocyathellus (Stapicyathus) stapipora (Taylor) Debrenne : 127.
1965 Robustocyathus stapipora (Taylor) Hill : 61 and 68.
LECTOTYPE. Taylor 1910, PI. 7, fig. 38 G, chosen here. The specimen should be
in the University of Adelaide.
OTHER MATERIAL. B.M. (N.H.) S 4733, S 4351-2, S 4138, S 4528, S 4817-20, S
7621.
DESCRIPTION. Narrow intervallum of constant width from the basal conical apex
to the upper bowl-shaped cup. Some specimens are more cylindrical. Outer wall
thin, pierced by round pores that are arranged quincunxially. Inner wall thicker,
with one vertical row of pores in front of each septa ; each pore is excavated into its
septum so that stirrup-pores are formed. These inner wall pores are also arranged
into horizontal lines.
DIMENSIONS
Cup:
Height (pars)
Diameter
Intervallum coeff.
Interseptum
Loculi
Outer wall:
No of pore rows per
interseptum
Diameter of pores
Vert, partitions
Thickness
Inner wall :
No. of pore rows per
interseptum
Diameter of pores
Vertical part.
Horizontal part
Thickness
Septa
Thickness
§4733
(mm.)
30
about 40
0-02
o-339
1/3-4
3
0-07
0-068
0-150
two 1/2
0-27
0-27
0-16
o-i
non porous
0-06
S 4351
(mm.)
40
about 20
0-45-0-66
0-61
from i/i -8 to 1/2
3
0-07
0-068
0-23
two 1/2
O'27
o-33
0-27
o-i
non porous
0-06
84352
(mm.)
S 4817-4820
(mm.)
22
about 12
15
about 35
o-i
0-61
1/2
0-042
0-74
1/2
3
0-06
0-07
3 to 4
0-075
0-068
0-15
o-i
tWO 1/2
tWO 1/2
O-2O
O-27
o-33
0-40
O-2O
O-27
O-I
0- I
non porous
non porous
0-07
0-07
S 7621
(mm.)
20
about 10
o-i
o-37
i/3'3
3
0-07
o-i
o- 1
tWO 1/2
0'27
O-27
O-2O
O-I
non porous
0-07
DISCUSSION. Only one species known. The different shapes i.e. cylindrical, or
bowl-shaped could conceivably represent different species, but their coefficients are
not sufficiently different.
310 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
Genus LOCULICYATHUS Vologdin, 1931
Subgenus LOCULICYATHELLUS nov.
TYPE SPECIES. Archaeocyathus floreus R. & W. R. Bedford 1934.
DIAGNOSIS. Cup has the generic characters of Loculicyathus : thin porous walls and
septa, vesicular tissue crossing intervallum and central cavity. The external longitu-
dinal corrugations on the outer wall, like those of Ajacicyathellus, subgenus of
Ajacicyathm (see Debrenne 1964 : 127), or Archaeocyathellus Ford 1873, distinguish it
as a subgenus.
DISCUSSION. R. & W. R. Bedford (1937) compared A. floreus with Archaeo-
cyathellus. They singled out forms with radial imperforate septa and stirrup-pores
at the inner wall, from others with regularly porous septa. The holotype of floreus
(S 4144), the paratype and other material in the B.M. (N.H.) collection, show sparse
pores but no stirrup-pores.
The only species recognized at present is Loculicyathus (Loculicyathellus} floreus
(R. & W. R. Bedford 1934).
Loculicyathus (Loculicyathellus} floreus (R. & W. R. Bedford)
(PI. 3, figs, i, 2, 4)
1934 Archaeocyathus floreus R. & W. R. Bedford : 2, fig. 4.
1937 Archaeocyathus floreus R. & W. R. Bedford; R. & J. Bedford : 35, figs. I44A & B.
HOLOTYPE. B.M. (N.H.) S 4144.
OTHER MATERIAL. Paratype B.M. (N.H.) S 4145. Also B.M. (N.H.) S 4730,
S 4739 and S 7635.
DIAGNOSIS. Small cylindrical cup with vertical corrugation between two neigh-
bouring septa, so that transverse sections recall scleractinian corolla. Outer wall with
horizontal and vertical rows of non-alternating pores. The pores of the inner wall and
septa are in quincunx. A few dissepiments occur across the intervallum and central
cavity.
DIMENSIONS
S 4144 S 4145 S 4739
(mm.) (mm.) (mm.)
Cup:
Height (pars) 9-5 10
flow 5 • o flow 5-0 6-77
Diameter < < "
\upp. 6-0 VUPP- 6'5
Interseptum o • 75-2 • o o • 75-2 • o 0-95
Interv. coeff. 0-4 .. 1-6
Parietal coeff. i -o . . i -6
Outer Wall:
No. of pore rows per intersept 6
Diameter of pores 0-13
Vertical skel. part. 0-13
Horizontal skel. part. °'33
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
DIMENSIONS — continued
Inner Wall :
No. of pore rows per intersept
Diameter of pores
Vertical skel. part.
Horizontal skel. part.
Septa :
Diameter of pores
Vertical skel. part.
Horizontal skel. part.
S 4144
(mm.)
2
0-23
o-33
0-23
0-23
o-33
0-23
s 4145
(mm.)
0-26-0 -37
o-35
O-22
0-30
0-26
S 4739
(mm.)
DISCUSSION. The specimens from Ardrossan figured by R. & J. Bedford, (1939,
figs. 144 A & B) only have one pore per intersept at the inner wall and a non-
corrugated outer wall. They would therefore, seem to be a quite different form.
Family ROBUSTOCYATHIDAE Debrenne, 1964
Genus ROBUSTOCYATHUS Zhuravleva, 1960
TYPE SPECIES. Archaeocyathus robustus Vologdin 1937, by original designation of
Zhuravleva (1960 : 133).
DIAGNOSIS. Solitary cups with simply porous outer wall and single vertical row of
apertures per intersept at the inner wall, so that each loculus opens into the central
cavity.
COMPOSITION OF THE GENUS. R. argentus (Okulitch 1935), R. annulatus Zhuravleva
1960, R. artecaveatus (Vologdin 19400), R. biohermicus Zhuravleva 1960, R. densus
Debrenne 1964, R. hupei Debrenne 1964, R. levigatus (Vologdin 19400), R. magnipora
(R. & W. R. Bedford 1934), R. moori (Vologdin 1937), R. novus Zhuravleva 1960, R.
polyseptatus (Vologdin 19400), R. proskurjakowi (von Toll 1889), R. pseudotichus
(Vologdin 19400), R. salebrosus (Vologdin 1931), R. spinosus Zhuravleva 1960,
R. subacutus (R. & W. R. Bedford, 1934), R. sucharichensis Zhuravleva 1960, R. ?
tennis (Vologdin 1940).
Robustocyathus magnipora (R. & W. R. Bedford)
(PI- 3, fig- 5)
1934 Archaeocyathus magnipora R. & W. R. Bedford : 2, pi. i, fig. 7.
1937 Paranacyathus magnipora (R. & W. R. Bedford) R. & J. Bedford : 34.
HOLOTYPE. B.M. (N.H.) S 4146.
DESCRIPTION. The holotype, a partially destroyed cup, was the only specimen
examined. The radial septa are few, straight and pierced by three alternate vertical
GEOL. 17, 7 25
312 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
rows of circular pores, which are not connected to the wall pores. The outer wall is a
thin plate with circular pores in quincunx. The inner wall has a single vertical row of
large pores per intersept and pores of neighbouring rows alternate. The septa, which
spring out of the inner wall, form boundaries to new pores by splitting the previous
one.
DIMENSIONS
Cup: (mm.)
Height (pars) 24
Diameter 8 • 8
Intervallum i • 83
Interseptum o • 74
Loculus (trapezoid) °'47 and 0-6 for
Height = 0-9
Parietal coefficient + unknown
Outer wall :
Vertical rows of pores per intersept 4
Diameter of pores o • 20
Skeletal partitions o • 13
Thickness 0-13
Inner wall : i
Vertical diameter o • 40
Horizontal diameter o • 60
Skeletal partitions 0-33
Thickness o • 33
Septa :
Alternating vertical rows of pores per septum 3
Diameter 0-61
Skeletal partitions 0-61
DISCUSSION. The specimens figured by R. & J. Bedford (1937, figs I42A-E) and
described as magnipora are unlike the holotype mentioned above. They differ by
having various forms and patterns of outer wall and septal pores. These characters,
together with the presence of an Archaeopharetra-type apex, caused R. & J. Bedford
to place magnipora in the genus Paranacyathus. Unfortunately, it was not possible
to compare the holotype with these specimens. The holotype (S 4146) shows the
true characters of Robustocyathus : regular walls and septal pores, which are sufficient
to place it in the Class Regularia, although the initial stages are not present in the
specimen. At the moment, it is not possible to decide the true systematic position
of the specimens figured by the Bedfords in 1937.
Robustocyathus subacutus (R. & W. R. Bedford)
(PL 3, figs. 6, 7, Text-fig. 4)
1934 Archaeocyathus subacutus R. & W. R. Bedford : 2, pi. i, figs. 3a-c.
1937 Archaeocyathus acutus Bornemann; Ting : 358, pi. 9, figs. 1-2.
1961 Ajacicyathus walliseri F. & M. Debrenne : 696, pi. 19, fig. 3.
LOWER CAMBRIAN ARCH AEOC Y ATM A FROM AUSTRALIA 313
HOLOTYPE. B.M. (N.H.) S 4142.
OTHER MATERIAL. B.M. (N.H.) S 4143, S 4747, S 4792, S 7620.
DESCRIPTION. Small cylindrico-conical cup with thin skeletal plates and few pores.
The outer wall only has one pore in the middle of each intersept; these pores are
arranged in widely separated vertical rows and more closely set horizontal ones. The
inner wall is thin and perforated by a single row of pores per intersept. The aperture
FIG. 4. Robustocyathus subacutus (R. & W. R. Bedford)
of each pore is larger than the width of the interseptum. The inner edges of the septa,
which laterally delineate the pore, have vertical undulations (see Text-fig. 4).
Numerous radial septa, having a few small isolated pores, are found near the outer
wall, but they have only been seen in specimen S 4747.
The undulations on the inner edges of the septa, close to the inner wall, are different
from those seen in Ethmophyllum Meek. The amplitude of the waves is smaller, while
the sinuous edges do not join together to close up the interseptum and form another
wall that connects the septa, but simply limit the inner wall pores. The inner wall,
itself, remains as a completely separate sheet.
3H LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
DIMENSIONS
S 4142 S 4143 S 4747 S 7620 walliseri
(mm.) (mm.) (mm.) (mm.) (mm.)
Cup:
Height (pars) 30 20 25 + +
Diameter 10-12 + 8-10 7 13
Intervallum coeff. 0-3 0-39 0-5 . . 0-23
Parietal coeff. 5-4 + 5-5 5-5 5-2
Interseptum 0-30 °'33 0-20 0-25
Loculus 1/6 1/8 1/7 . . 1/4
Outer wall:
No. of pore rows per intersept i + + i i
Diameter 0-13 .. .. 0-7 0-13
Skeletal vert. part. 0-40 .. .. 0-40 +
Skeletal horiz. part. 0-13 .. .. 0-13 o-n
Thickness 0-06 . . 0-06 0-07 0-07
Inner wall :
No. of pore rows per intersept i i i i i
Diameter 0-33 0-40 0-27 0-27 0-26
Skeletal vert. part. 0-13 0-13 + 0-13 0-13
Skeletal horiz. part. 0-13 0-13 + 0-13 +
Thickness 0-33 0-40 0-33 0-27 0-26
Septa
Thickness o-io o-io + o-io 0-06
DISCUSSION. R. walliseri Debrenne, in spite of its very large size, seems to fall
between the limits of specimens of subacutus. Another species, R. pseudotichus
(Vologdin) has similar undulations on the inner parts of the septa that border the
large inner wall pores, but in addition, has long spines on the vertical edges of the
pores.
Genus ZONACYATHUS R. & J. Bedford, 1937
1937 Zonacyathus R. & J. Bedford : 36.
I94oa Ethmophyllum Meek; Vologdin (pars) : 66-68.
TYPE SPECIES. Archaeocyathus retevallum R. & W. R. Bedford (1934 : 2, fig. 6), by
monotypy.
DIAGNOSIS. Porous two-walled cups with no, or few, porous radial septa. The
outer wall has regular pores. The inner wall has branching pore-tubes; the initial
tube is located in the middle of each interseptum and then branches so that the
secondary tubes open in front of the septa. The tubes may lengthen and curve into
the central cavity. The pore-tubes are never formed by septal fluting.
DISCUSSION. The type species retevallum is known from only the three specimens
in the B.M. (N.H.) collection. The skeletal tissue is extremely thin and breakable,
with the result that structures are either crushed or destroyed, making it difficult to
distinguish the pore systems. The validity of a genus based on such fragments may
be questionable. Nevertheless, from studying a very close but simpler species, Z.
retezona Taylor, which has an inner wall built of shorter pore-tubes that branch in the
same way, it is possible to understand the more complex pattern present in retevallum.
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 315
COMPOSITION OF THE GENUS. Zonacyathus retevallum (R. & W. R. Bedford 1934),
Z. retezona (Taylor 1910). The species Ethmophyllum poletevae Vologdin 19400;, E.
vermiculatum Vologdin 1938 and E.flexum Vologdin 1940^, are placed in Zonacyathus
with reservation.
Zonacyathus retevallum (R. & W. R. Bedford)
(PL 4, figs. 4, 5)
1934 Archaeocyathus retevallum R. & W. R. Bedford : 2, fig. 6.
1937 Zonacyathus retevallum (R. & W. R. Bedford,) R. & J. Bedford : 36, figs. I53A-E.
1965 Zonacyathus retevallum (R. & W. R. Bedford): Hill : 76, pi. 4, figs. 3a-b.
HOLOTYPE. B.M. (N.H.) S 4147.
OTHER MATERIAL. B.M. (N.H.) S 4726-4727.
DESCRIPTION. Bowl-shaped cup with narrow waved intervallum, easily distorted
but without giving any obvious folds. Owing to the bad preservation of the material,
all measurements are approximate and observations incomplete.
Outer wall simple with two alternating rows of oval pores, arranged in quincunx, to
each interseptum. The septa are difficult to see in longitudinal section in the Bed-
ford specimens. The septa appear to be imperf orate, but may have small scattered
pores. The inner wall has branching pore-tubes of Y-form, which lengthen and curve
into the central cavity. It is suspected that some lateral communication occurs
between the pore-tubes that penetrate into the central cavity.
DIMENSIONS
S 4147 : i S 4147 : 2 S 4726-4727
(mm.) (mm.) (mm.)
Cup:
Height 25 40
Diameter 20 12-30 10
Intervallum coefficient o • 1 1 0-12 0-2
Parietal coefficient . . 6-6
Interseptum 0-27 0-27 0-27
Loculus J/7'2 J/4'8 I/5'5
Outer wall:
No. of rows of pores per intersept 2 2
Diameter of pores .. 0-13X0-06
Vertical partitions .. o-io
Horizontal partitions .. o-io
Thickness o-io 0-13 o-io
Inner wall :
No. of rows of pores per intersept i i i
Diameter of pores 0-20 0-27 0-20
Vertical partitions o-io 0-13 0-13
Horizontal partitions o-io 0-13 0-13
Thickness at least 0-6 0-8 0-6
Septa :
Thickness o • i o o • i o
3i6
LOWER CAMBRIAN A RCH AEOC Y ATM A FROM AUSTRALIA
DISCUSSION. Ethmophyllum flexum Vologdin (19400 : 66, pi. 15, fig. 4) is very
similar to Z. retevallum R. & W. R., Bedford, having the same irregular shape and
size.
Zonacyathus retezona (Taylor)
(PI. 4, figs, i, 2, 3, Text-fig. 5)
1910 Archaeocyathus retezona (Taylor) : 121, pi. 7, fig. 38A, pi. 6, fig. 3iE.
MATERIAL. B.M. (N.H.) S 4353, S 4764, S 4774, S 4778, S 4803-4808, S 4811-4816,
S 7623, S 7634, S 4731.
DESCRIPTION. Conical, externally smooth cups. The outer wall is a porous sheet,
with vertical alternating rows of slightly elliptical pores surrounded by skeletal tissue
FIG. 5. Zonacyathus retezona (Taylor)
of constant width. Radial septa are, probably, imperforate. Inner wall independent
from septa and has two vertical rows of pores, one row opposite each septa and the
other in the middle of each interseptum. Each pore opposite a septum, is the opening
for two oblique pore-tubes, one coming from each loculus. On the inner face of the
intervallum, a central, radial pore-tube is joined by a branch from each of its neigh-
bours to form one elliptical pore. All of these pore-tubes are horizontal. Z. retezona
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
317
differs from Z. retevallum because its tubes are short and do not lengthen, or curve into
the central cavity.
DIMENSIONS
S 4811- S 4730-
S 4774-
Cup:
Height
Diameter
Intervallum coeff.
Loculus
Parietal coeff.
Outer wall :
No. or rows of pores
per intersept
Diameter of pores
Vertical partitions
Horizontal partitions
Thickness
Inner wall :
No. of rows of pores
per intersept
Diameter of pores (
Vertical partitions
Horizontal partitions
Thickness
Septa :
No. of pore rows
Diameter of pores
Partitions
Thickness
S4353
(mm.)
20
10
O-2I
3'9
4
0-06
0-07
0-06
0-13
87623
(mm.)
8
O-22
4-2
0-13
4816
(mm.)
45
15
O-I2
3
o-io
0-07
0-06
0-13
4732
(mm.)
42
27
0-9
4
0-06
0-06
0-07
0-13
S 4764
(mm.)
4777
(mm.)
S7634
(mm.)
84778
(mm.)
20
pars 15
. .
13
5-8
9
10
10
1/2
4-8 to 3*9
0-25
O-2I
3'4
O-2I
4-6
0-07
0-06
0-06
o-io
I
i
i
i
I
20-0-27
0-27
0-13-0 -2
0-27
0-27
0-16
0-13
O-IO
0-13
0-13
0-13
0-13
O-2O
O-20
0-40
0-40
o-35
0-40
0-40
4-6 4-6
0-13
I
0-27
0-16
0-54
4
0-07
0-05
I
O-2O
0-13
0-40
0-07
O-IO
2
0-13
0-13
O-IO
0-13
O-IO
0-06 0-07
DISCUSSION. Taylor (1910 : 122) gave the following interpretation of the inner
wall of Z. retezona: " a simply porous inner wall, with one or two pores between each
pair of adjacent septa ". In fact, if one refers to his fig. 27, it is seen that one pore
occurs in each septum and the other in the middle of the interseptum. I consider
that the regular inner wall is composed of pore tubes, that join together and open
into the central cavity. Taylor thought the inner parts of the septa were modified
into curved rods which supported the simple inner wall. However, the excellent
silicified material in the Bedford collection enables the path of the canals to be
followed throughout their length.
Family BRONCHOCYATHIDAE R. & J. Bedford, 1936
1936 Bronchocyathidae R. & J. Bedford : 25
1937 Stillicidocyathidae Ting : 367.
1951 Thalamocyathidae Zhuravleva : 98.
1955 Ethmophyllidae Okulitch : E 12.
J959 Cyclocyathellidae Zhuravleva : 426.
1965 Bronchocyathidae R. & J. Bedford; Hill : 93.
TYPE GENUS.
1936).
Thalamocyathus Gordon 1920 (=Bronchocyathus R. & J. Bedford,
3i8 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
COMPOSITION of GENUS. Thalamocyathus Gordon 1920, Stillicidocyathus Ting 1937.
Polystillicidocyathus Debrenne 1959, Cyathocricus gen. nov.
REMARKS. Thalamocyathidae Zhuravleva 1951 has not won general acceptance
under Article 400 of the International Code of Zoological Nomenclature, 1961 and
while this rule stands in its present form, this family name must be rejected.
Genus CYATHOCRICUS nov.
TYPE SPECIES. Archaeocyathus tracheodentatus R. & W. R. Bedford, 1934.
DERIVATION. From KPIKOS, cricos = a ring.
DIAGNOSIS. Cup with a simply porous outer wall; straight, sparsely perforated
radial septa and an annulate inner wall. Annuli consist of undulating, horizontal, or
slightly inclined plates that are neither S-, nor V-shaped ; their axial rim is cogged.
COMPOSITION OF THE GENUS. C. tracheodentatus (R. & W. R. Bedford 1934), C.
dentatus (Taylor 1910) and C. annulispinosus (Vologdin 1931).
DISCUSSION. Cyathocricus differs from other annulate genera in that the rings
appear to be straight in vertical section. The ft component, closest to the septa, is
generally horizontal and the a component, projecting into the central cavity, is
complicated by cogs that are more or less fused together.
This new genus is established for those species incorrectly referred to the genus
Bronchocyathus R. & J. Bedford 1936. Hill (1965 : 94) gives an account of this
situation, involving Thalamocyathus and Bronchocyathus.
Gordon (1920 : 687) created the genus Thalamocyathus and included the species
Archaeocyathus tubavallum Taylor, A. trachealis Taylor, A. infundibulum Bornemann,
A . ichnusae Meneghini and T. flexuosus Gordon, but did not designate a type species.
R. & J. Bedford (1936 : 25) erected Bronchocyathus, designating B. trachealis
(Taylor) as the type species and including Ethmophyllum dentatum Taylor and
Archaeocyathus tracheodentatus R. & W. R. Bedford.
The type species of Thalamocyathus Gordon was subsequently designated by Ting
(J937 : 368) as T. trachealis (Taylor), after elimination of the other syntype species
because they possessed cribrose inner walls. Hill (1965 : 94) states that she con-
siders this to be a valid designation. Simon (30th Dec. 1939 : 40) later named A.
trachealis as the type species of the genus, in case of any doubts that Ting had done so.
Thus Bronchocyathus R. & J. Bedford, is a junior objective synonym of Thalamo-
cyathus Gordon since both have the same type species (Int. Code Nomen., Article 6ib).
R. & J. Bedford (1939 : 75) pointed out that A. trachealis Taylor belonged to
Cyclocyathus Vologdin 1931, a genus of which they were unaware in 1936. As
Vologdin had not designated a type species, they cited trachealis, but this is not
acceptable since the species was not listed in the original description of Cyclocyathus,
although the Bedfords considered that Vologdin's use of tubavallum Taylor was a
lapsus calami for trachealis Taylor. In referring to tubavallum, the Bedfords
mentioned that it was the first species listed by Gordon in his description of Thalamo-
cyathus and was founded on a single fragment of which conflicting figures were pro-
vided, and that the whereabouts of the specimen is uncertain. Finally, R. & J.
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA 319
Bedford (1939 : 75) decided to apply the name Bronchocyathus to those forms having
a more complex inner wall than Cyclocyathus and gave Ethmophyllum dentatum
Taylor as the type species. This does not conform to the rules of zoological nomen-
clature, for Article 61 states "the type of any taxon, once fixed in conformity with the
provisions of the Code, is not subject to any change except by exercise of the plenary
powers of the Commission ". It is, therefore, necessary to provide another name for
the genus containing dentatum Taylor. Archaeocyathus tracheodentatus R. & J.
Bedford, is selected as the type species of the new genus Cyathocricus, as it is more
thoroughly known than dentatum.
Hill (1965 : 94) mentions other nomenclatural problems of Thalamocyathus and the
species trachealis Taylor. Zhuravleva (1960 : 220) cites Bronchocyathus as the genus
of the family Bronchocyathidae, distinguishing this group from the Stillicidocyathidae
by the presence of inter-communicating pore-tubes. However, this was based on the
use of E. dentatum Taylor as type-species of Bronchocyathus.
Cyathocricus tracheodentatus (R. & W. R. Bedford)
(PL 5, figs. 4, 5, PL 6, fig. 4, Text-fig. 6)
1934 Archaeocyathus tracheodentatus R. & W. R. Bedford : 2, fig. 5.
1936 Bronchocyathus tracheodentatus (R. & W. R. Bedford) R. & J. Bedford : 25, fig. 104.
HOLOTYPE. B.M. (N.H.) S 4148.
OTHER MATERIAL. B.M. (N.H.) S 4754.
DESCRIPTION. The type specimen is a conical cup with a broken apex. Corruga-
tions on the upper part provide good tangential and longitudinal sections of the inner
wall (see Text-fig. 6) . The numerous radial septa are straight and are perforated by a
few vertical rows of fine pores. Part of the outer wall is preserved, showing the some-
what irregularly placed polygonal pores. The inner wall consists of a vertical series
of thick, horizontal, ring-shaped plates, which are triangular in cross-section and are
joined to the inner edges of the septa by the /? component. The annuli thin toward
the central cavity, are slightly wavy on their upper face and their free rim (a com-
ponent) is regularly cogged.
DIMENSIONS
S 4148 (Upper) S 4148 (Lower) S 4754-4755
(mm.) (mm.) (mm.)
Cup:
Height (pars). 31 31 5
Upper Diameter 1 1 6-5 7
Lower Diameter 0-2 0-3 0-3
Parietal coefficient . . 8-3
Interseptum 0-27 0-13 0-13
Loculus 0-27/1-6 0-13/1-29
Outer wall :
No. of pore rows per intersept 3-4 . . 2
Diameter 0-06 0-06 0-05
Vertical partitions 0-05 0-05 0-05
Horizontal partitions 0-05 0-05 0-05
Thickness o-io o-io 0-13
32°
LOWER CAMBRIAN ARCH AEOC Y ATM A FROM AUSTRALIA
DIMENSIONS — continued
Inner wall:
Space between rings
Thickness of ring
Length of cog
Width of ring
Space between cogs
Septa :
No. of pores
Thickness
S 4148 (Upper) S 4148 (Lower)
(mm.)
0-27
0-13-0-20
0-67
o-33
Imperforate
0-06
4754-4755
(mm.)
O-2O
O-IO
0-67
O-27
Imperforate
0-05
DISCUSSION. C. annulispinosus (Vologdin 1931) is similar morphologically, but its
specific coefficients are not well known. According to Vologdin (1931 : fig. 42) the
diameter is approximately 5-6 mm, the intervallum coefficient would be 0-4 and the
parietal coefficient 12 — figures that are quite close to those of specimen S 4148. The
essential difference is in the downward orientation of the annuli into the central
cavity present in annulispinosus.
FIG. 6. Cyathocricus tracheodentatus (R. & W. R. Bedford)
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 321
Cyathocricus dentatus (Taylor)
(PI. 6, figs. 1-3, Text-fig. 7)
1910 Ethmophyllum dentatum Taylor : 129, pi. 10, fig. 59; pi. 13, fig. 76; pi. 16, fig. 89; t.-figs.
23 and 31.
1936 Bronchocyathus dentatus (Taylor) R. & J. Bedford : 25, fig. 103.
1960 Bronchocyathus dentatus (Taylor); F. & M. Debrenne : 703, pi. 20, fig. 8.
HOLOTYPE. Not designated.
OTHER MATERIAL. B.M. (N.H.) S 4752-4753, S 4756, S 4355.
DESCRIPTION. Cylindrical-conical cup that may have a large diameter and a
wide central cavity. The intervallum is full of straight radial septa, that have about
2-3 vertical rows of small pores near the outer wall. The outer wall is perforated by
pores of the same size as those of the septa, with 2-3 alternate vertical rows per
interseptum. There are no stirrup-pores connecting the two systems. The inner wall
is formed by a series of vertical annular plates, which are as thick as the septa. These
plates extend into the inner part of the intervallum, as well as into the central cavity.
FIG. 7. Cyathocricus dentatus (Taylor)
322 LOWER CAMBRIAN ARCH A EOC Y ATM A FROM AUSTRALIA
Their free edge is serrated into toothlike projections, that are 2-3 loculi wide at the
base, and which are situated either in front of a septum, or in the centre of an inter-
septum (PI. 6, fig. 2). The tips of these projections are flat on top and bend slightly
upwards. When examining the material, layers of the teeth were prised apart without
discovering any order of appearance. On the lower part of each tooth there is a curved
support which joins it to the underlying ring plate.
DIMENSIONS
S4355
(mm.)
S4753
(mm.)
12
o-3
0-13
2
O-O6
0-05
0-05
O-O6
30
18
0-15
O-2O
2-3
O-O6
0-05
O-O5
O-O6
S4756
(mm.)
53
15-16
S 4752
(mm.)
9
0-13
not seen
0-06
0-13
Cup:
Height
Diameter
Intervallum coefficient
Interseptum
Outer wall:
No. of pore rows per intersept
Diameter
Vertical partitions
Horizontal partitions
Thickness
Inner wall:
Distance between annuli
Thickness of annuli
Width of annuli
Projections
Septa :
Thickness
Diameter
Vertical partitions
Horizontal partitions
DISCUSSION. The interpretation of the inner wall structure of dentatus is difficult
owing to the limited number of known specimens. In common with other forms, the
term " vesicular " has been used in the past to describe the various poor oblique
sections of specimens, which have a complex inner wall structure. It was not
possible to examine the holotype, but there are specimens of dentatus in the Bedford
collection, wrongly called tracheodentatus, which provided various views of this
structure. From these specimens (S 4355, S 4753 and S 4752) it was possible for me
to reconstruct the inner wall of dentatus with some certainty. This species differs
from tracheodentatus and annulispinosus by having thinner annuli, vertical supports
between the annuli, and the triangular shape and different distribution of the teeth.
0-20
O-O6
0-27
0-06
0-20
0-06
O-2O
o- 10
0-40
0-40
Imperforate
o-33
0-67
Perforate
on outer side
0-27
0-27
Imperforate
o-33
0-67
0-06
0-06
0-06
0-05
0-06
0-27
Family ETHMOCYATHIDAE nov.
DIAGNOSIS. Simple outer wall; uniserially pored inner wall covered over on the
innermost (central cavity) side with a secondary wall of annular plates. This feature
is characteristic of the new family. Ethmocyathus R. & W. R. Bedford is the type
genus.
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA 323
DISCUSSION. It is necessary to create a separate family within the Ajacicyathacea
to place the genus Ethmocyathus. There is no justification for placing Ethmocyathus
in the family Ethmophyllidae (Hill 1965 : 76; Zhuravleva 1960 : 162), for its inner
wall is not formed by horizontal fluting of the inner edges of the septa, as they
suggested. Neither is the genus a doubtful member of the Tumulocyathidae
(Debrenne 1964 : 113), since the outer wall has simple pores and its annuli have a
peculiar form.
Genus ETHMOCYATHUS R. & W. R. Bedford, 1934
1934 Ethmocyathus R. & W. R. Bedford : 2, fig. 8.
TYPE SPECIES. Ethmocyathus lineatus R. & W. R. Bedford, by monotypy.
DESCRIPTION. Cup with straight radial, sparsely porous septa. The outer wall
has close, simple pores. The inner wall is composed of a thin sheet of hexagonal pores
screened from the central cavity by thin, narrow horizontal annuli.
DISCUSSION. Sometimes, both of the horizontal sides of the inner wall hexagonal
pores are reduced so that the openings become rhombic. The inner edges of the
septa are sinuous, following the outlines of the inner wall pores, but do not contribute
to their formation by horizontal fluting.
COMPOSITION OF THE GENUS. Ethmocyathus lineatus R. & W. R. Bedford.
Ethmocyathus lineatus R. & W. R. Bedford
(PI. 5, figs., 1-3, Text-fig. 8)
1934 Ethmocyathus lineatus R. & W .R. Bedford : 2, fig. 8.
1965 Ethmocyathus lineatus R. & W. R. Bedford; Hill : 76, pi. 4, figs. 2a-b, t.-fig. 16. 2.
HOLOTYPE. B.M. (N.H.) S 4149.
DESCRIPTION. This is based on a single specimen consisting of a fragment from a,
probably, cylindrical cup. Radial septa with sparse pores on the outer two-thirds of
each septum. The annular plates join the inner wall net of hexagonal pores, without
any visible thickening. The outer wall is simple with regular oval pores.
DIMENSIONS
Cup: (mm.)
Height (pars). 15
Diameter (approx.) 13
Interseptum o • 1 8
Intervallum coefficient 0-13
Outer wall : (only seen between 2 neighbouring septa)
3 pores lengthened horizontally and arranged in quincunx
Diameter o • 05-0 • 03
Skeletal partitions o • 03
Thickness o • 07
LOWER CAMBRIAN ARCHAEOCYATHA FROM AUSTRALIA
DIMENSIONS — continued
Inner wall:
Vertical diameter
Horizontal diameter
Skeletal partitions
Thickness
Thickness of annuli
Distance between annuli
5 annular plates cover one pore (vertically)
Septa:
Imperf orate for 0-37 mm. from the inner wall
Diameter of pores
Vertical partitions
Horizontal partitions
DISCUSSION. Only this one species known.
(mm.)
0-27
0-2
0-075
o-37
0-03
0-03
0-03
0-15
0-18
FIG. 8. Ethmocyathus lineatus R. & W. R. Bedford
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA 325
Family ERISMACOSCINIDAE Debrenne, 1964
DIAGNOSIS. Cup with radial septa and porous tabulae. Thin, simply porous
outer wall. Inner wall with several vertical series of pores to each interseptum,
sometimes with thickening of the pore framework (" linteaux ") and ornamental
spines.
COMPOSITION OF THE FAMILY. Erismacoscinus (Pluralicoscinus) Debrenne 1963,
Erismacoscinus Debrenne 1958, Asterocyathus Vologdin 1940, Retecoscinus Zhurav-
leva 1960, Tuvacyathus Vologdin 1940, Geniculicyathus Debrenne 1960, Coscinoteichus
Debrenne 1964, Coscinoptycta Broili 1915.
Genus ERISMACOSCINUS Debrenne, 1958
TYPE SPECIES. Erismacoscinus marocanus Debrenne 1958 : 65, pi. 3, figs. 11-16,
by monotypy.
DIAGNOSIS. Intervallum with regularly porous septa and tabulae. Simple outer
wall, generally perforated by numerous openings in quincunx. Thick inner wall,
always has several vertical series of pores to each interseptum. The inner wall pores
frequently carry spines, that vary in shape and distribution.
COMPOSITION OF THE GENUS. Erismacoscinus contains all those species, having
several vertical series of pores to each interseptum, that were previously included in
the genus Coscinocyathus Bornemann (see list of Debrenne 1964 : 166-167).
DISCUSSION. Hill (1965 : 108-109) suggests this genus might be synonymous with
Tannuolacyathus Vologdin 1957, but the vesicular tissue and double porous wall of
the latter, have led me to consider that genus as belonging to a distinct family, the
Tannuolacyathidae (Debrenne 1964 : 188) .
Erismacoscinus rugosus (R. & W. R. Bedford)
(PI. 8, figs. 2, 3)
1934 Coscinocyathus rugosus R. & W. R. Bedford : 3, fig. n.
1964 Erismacoscinus rugosus (R. & W. R. Bedford) Debrenne : 167.
HOLOTYPE. B.M. (N.H.) S 4152.
DESCRIPTION. A single specimen of a funnel-shaped cup with a wide intervallum
and rather narrow central cavity. Septa simply porous and not always reaching
the inner wall. Tabulae form a regular network of pores. The inner wall is simple
with some spines. The pores of the outer wall are in vertical rows, between which the
" linteaux " project. These keels continue towards the exterior, giving the fossil a
rough appearance from which its specific name is derived. The pores of neighbouring
rows alternate slightly.
326 LOWER CAMBRIAN ARCH AEOCYATH A FROM AUSTRALIA
DIMENSIONS
(mm.)
Cup:
Height (pars). 6
Diameter from 4 • 75-5 • 5
Interseptum (outer) i-oi
(inner) 0-33
Intertabulum I • 35
Loculi 1/2, 8/2, 8
Intervallum coefficient 0-8
Parietal coefficient 3 • 6
Outer wall :
2-3 rows of pores per interseptum
Vertical diameter of pores o • 20
Horizontal diameter of pores o • 30
Vertical thickness 0-13
Horizontal thickness o • 05
Inner wall:
2 rows of alternating pores per interseptum
Diameter of pores o • 10
Linteaux 0-16
Thickness of wall o-io
Septa :
10 rows of pores per septum
Diameter of pores o • 05
Partitions 0-15
Tabulae :
7 pores per interseptum
Diameter of pores o • 05
Skeletal partitions 0-06
DISCUSSION. The presence of outer keels is a rare modification. It seems that
E. equivallum (Taylor) has outer keels in front of each septum (see below).
Erismacoscinus equivallum (Taylor)
(PI. 7, figs. 1-3 and 5)
1908 "turbinate Archaeocyathinoid" Taylor : 426, pi. i, fig. i.
1910 Coscinocyathus equivallum Taylor : 138-139, pi. 15, fig. 85, t.-fig. 12.
1964 Erismacoscinus equivallum (Taylor) Debrenne : 167
HOLOTYPE. In the University of Adelaide.
OTHER MATERIAL. B.M. (N.H.) S 200 is the counterpart (i.e. the other face of the
section) of the holotype. S 198, naturally etched specimen figured by Taylor (1908,
pi. i, fig. 2 : 423).
DESCRIPTION. Outer wall has projections of skeletal tissue corresponding to the
septa. As I did not see a specimen entirely free of matrix, it is only possible to
assume that this feature is similar to the keels present in E. rugosus. These projec-
LOWER CAMBRIAN ARCH AEOCY ATM A FROM AUSTRALIA 327
tions only occur at the junction of the septa and the outer wall, and not between the
rows of pores in the interseptum. The pores are elliptical and in quincunx.
The inner wall has round, alternate pores. The linteaux are smooth on the inter-
vallum side but small projections of spines and " bristles " occur on the central cavity
face.
Thick, straight septa have round regularly scattered pores in elongated quincunx.
The skeletal tissue occupies a greater surface area than the pores. Slightly convex
tabulae have a fine network of small, more or less regular polygonal pores.
DIMENSIONS
S 200 S 198
(mm.) (mm.)
Cup:
Height 22
Diameter 10 30-18
Intervallum coefficient 0-46 °'43
Parietal coefficient Not measurable 66
Interseptum o • 3 7 0-41
Intertabulum o • 44
Outer wall:
No. of pore rows per intersept 2 2
Diameter of pores 0-15 0-22-0-15
Vertical partitions o-n 0-11-0-15
Horizontal partitions 0-15 o • 1 1
Thickness o • 1 1 o • 1 1
Inner wall :
No. of pore rows per intersept 2 2
Diameter of pores 0-07 0-15
Vertical partitions 0-07 o • 1 1
Horizontal partitions 0-07 o • 1 1
Thickness o • 1 1 o • 1 1
Septa :
No. of pore rows per intersept 9-10 15
Diameter of pores o • 1 1 0-07
Vertical partitions 0-15 0-22
Horizontal partitions 0-15 0-22
Thickness 0-07 0-15
Tabulae:
No. of pore rows per intersept 14 4-22
Diameter of pores 0-05 o • 05-0 • 15
Vertical partitions 0-05 o • 03-0 • 07
Horizontal partitions 0-05
Thickness 0-07 0-07
DISCUSSION. Specimen S 198 is somewhat corrugated. At the level of the folds
the septa are not radial but parallel, as seen in Polystillicidocyathus Debrenne and
other colonial forms.
The similarity in structure of the two walls is not quite so evident as Taylor stated
(1910 : 138). The specimen S 200 shows that the inner wall is a little thinner and has
ragged " linteaux " and spines.
GEOL. 17, 7 26
328 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
Erismacoscinus quadratus (R. & W. R. Bedford)
(PI. 7, fig- 4> PL 8, fig- 5)
1934 Coscinocyathus quadratus R. & W. R. Bedford : 3, pi. 2, fig. 10.
1964 Erismacoscinus quadratus (R. & W. R. Bedford) Debrenne : 167.
HOLOTYPE. B.M. (N.H.) S 4157.
DESCRIPTION. Only one incomplete specimen known; a little deformed and there-
fore, difficult to measure accurately or to calculate the specific coefficients.
The septa and tabulae form a sub-quadrate network of irregular shape, due to the
slight waving of the intervallum plates. They both have isolated, simple, round
pores arranged in quincunx. The outer wall also has simple pores in quincunx but
with the majority often lengthened horizontally. The inner wall is smooth on the
intervallum side, but irregular and with spines around the pores on the central cavity
face. The pores are arranged in horizontal rows and alternate from one row to
another.
DIMENSIONS
Lower section Upper section
(mm.) (mm.)
Cup:
Height 25
Diameter n 16
Intervallum 2-64 2-64
Interseptum 0-37-0 -50 Q'37
Intertabulum o • 30-0 -60
Intervallum coefficient 0-38 0-2
Parietal coefficient probably 3-1
Outer wall :
3-4 rows of pores per intersept
Diameter of pores o • 07-0 -15
Partitions o • 1 1
Thickness 0-05
Inner wall:
2-3 rows of pores per intersept
Diameter of pores o • i i-o • 13
Septa :
20 pores per loculus
Diameter of pores 0-03
Partitions o • 1 1
Thickness 0-05
Tabulae :
3 rows of 20 pores per loculus
Diameter of pores 0-03
Partitions 0-06
Thickness o- 15 (with the spines)
DISCUSSION. The peculiar features of the inner wall (smooth in the intervallum.
spined in the central cavity) have previously been described for E. rugosus (R. & W. R.
Bedford) and E. equivallum (Taylor) — two species that differ from quadratus in other
characters.
LOWER CAMBRIAN ARCHAEOCYATHA FROM AUSTRALIA 329
Okulitch (1948 : 343) compared rhyacoensis and quadmtus, as both have subquad-
rate loculi. However, the American species has pore-tubes in the outer wall; an
inner wall which is typical of Coscinocyathus, with a single pore series to each inter-
septum; and an inter vallum with vesicular tissue as well as tabulae.
Erismacoscinus textilis (R. & W. R. Bedford)
(PI. 9, figs. 1-4)
1934 Coscinocyathus textilis R. & W. R. Bedford : 3, pi. 2, below fig. n.
1964 Erismacoscinus textilis (R. & W. R. Bedford) Debrenne : 167.
HOLOTYPE. B.M. (N.H.) S 4155.
PARATYPE. B.M. (N.H.) S 4156.
DESCRIPTION. The holotype has a cylindro-conical cap that tapers gently in its
lower parts. The intervallum consists of quadrate to rectangular loculi, limited by
radial septa and flat, irregularly-spaced tabulae.
The outer wall has a single vertical series of pores per interseptum (in a large inter-
septum there are two rows, but these soon become separated by a new septum). The
pores of the outer wall are much larger than those of the inner wall, they are also
horizontally lengthened and the rows alternate. Pores of the inner wall are very
small and arranged in quincunx. Septa and tabulae are regularly porous. Skeletal
tissue is well-developed.
The paratype is a fragment of a large, bell-shaped cup, with a narrow intervallum
in which the different pore characters of the two walls can easily be compared. The
inner wall has a regular net of round pores in quincunx, whilst the outer wall has 1-2
rows of pores. Domes of vesicular tissue are present in several loculi.
DIMENSIONS
S 4155 (Upper) S 4155 (Lower) S 4156
(mm.) (mm.) (mm.)
Cup:
Height : 30 43
Diameter 10 6 36 (Chord)
Intervallum 1-32 1-32 2
Interseptum 0-40 °'35 0-62
Intertabulum 0-6-2-71 irregular 2-70-4-70
Parietal coefficient 4-2 3-3
Outer wall :
No. of rows of pores per interseptum 1-2 1-2 1-2
Diameter 0-34-0-15 0-34-0-15 0-15
Skeletal partitions o-n o-n 0-15
Horizontal partitions 0-07 .. 0-15
Thickness 0-95 .. 0-15
Inner wall :
No. of pore rows per interseptum 3-4 . . 4
Diameter 0-07 .. o-io
Vertical partitions 0-07 .. o-io
Horizontal partitions 0-07 .. o-io
Thickness o • n . 0-07
330 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
DIMENSIONS — continued
S 4155 (Upper) S 4155 (Lower) S 4156
(mm.) (mm.) (mm.)
Septa :
No. of rows of pores 8 . . 10
Diameter o-n .. o-io
Vertical partitions 0-15 .. o-io
Thickness 0-07 .. o-io
Tabulae :
No. of pores per interseptum 4-5 4 5
Diameter 0-07 0-07 0-05
Vertical partitions 0-07 0-07 0-07
Horizontal partitions . . . . 0-07
Thickness 0-05 0-05 o • i o
Vesicular tissue : . . . . 0-03
DISCUSSION. The two specimens, a cylindrical one and a bowl-shaped one, are
placed in the same species. More material is needed in order to decide whether they
belong to separate species, or not. Generally the outer wall of Archaeocyatha is
thinner than the inner one. However, in this species the opposite is the case. It is
impossible to recognize the correct position of a fragment, if one only has the pore
characters of a wall to go by. Another example of this difficulty is the case of
Coscinocyathus unilinearis R. & W. R. Bedford (see PI. 13, fig. 4), and Coscinoptycta
convoluta (Taylor). If the wall with a single pore, is the inner wall, then Coscino-
ptycta is a synonym of Coscinocyathus and unilinearis is a species of Coscinocyathus.
On the other hand, if it is the outer wall that has one pore, then Coscinoptycta remains
a separate genus and unilinearis is a species of Erismacoscinus.
Erismacoscinus cellularis (R. & W. R. Bedford)
(PI. 8, figs, i and 4)
1934 Coscinocyathus cellularis R. & W. R. Bedford : 3, pi. 3, fig. 16.
HOLOTYPE. S 4162 B.M. (N.H.).
DESCRIPTION. The holotype is a longitudinally broken fragment of an almost
cylindrical cup, showing the porosity of the different skeletal plates.
The outer wall is a thin, regular and finely porous plate with horizontally elongated
pores. The inner wall is thicker, consisting of small, regular polygonal tubes that face
upwards; its central cavity face has several spines, or other outgrowths, arising from
the pore walls.
The septa have small evenly-spaced pores. The tabulae are close together with
their convex side uppermost, they are also finely perforated but the pores are more
numerous than those of the septa.
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 331
DIMENSIONS
(mm.)
Cup:
Height 55
Diameter 20
Intervallum 3
Intertabulum 1-1-32
Interseptum o • 30
Intervallum coefficient o • 75
Parietal coefficient not measurable
Outer wall :
No. of pore rows per interseptum 2
Diameter o • 1 5-0 • 07
Vertical partitions o • 07
Horizontal partitions o • 1 1
Thickness o • 07
Inner wall :
No. of pore rows per interseptum 3
Diameter 0-15; o-u ; o-n (nr.
central cavity)
Vertical partitions o • 1 1
Thickness 0-26
Septa :
No. of pore rows per interseptum 30 approx.
Diameter o • 03
Vertical partitions o • 1 1
Horizontal partitions o • 08
Thickness o • 07
Tabulae :
No. of pore rows per interseptum 3-4
Diameter o • 05
Vertical partitions o • 07
Horizontal partitions o • 07
Thickness o • 07
DISCUSSION. R. & W. R. Bedford (1934 : 3) suggested that the outer wall is
covered by a thin membrane that has very minute pores, but I could not find any
evidence of this structure. Their " irregular mosaic " of the inner wall is, in fact, the
result of the modification of the skeletal tissue (" linteaux ") into barbs and spines.
The regular polygonal tubes can be seen on the intervallum face of the inner wall,
as in E. rugosus and E. equivallum.
Erismacoscinus peter si (R. & W. R. Bedford)
(PI. 10, figs. 4 and 5)
1934 Coscinocyathus petersi R. & W. R. Bedford : 3, pi. 3, fig. 13.
HOLOTYPE. B.M. (N.H.) S 4158.
DESCRIPTION. The holotype is a unique, small, well-preserved specimen. The
outer wall is simply porous with prominent, sharply-ridged, vertical crests between
the septa. The inner wall is a skeletal plate that is perforated by vertical and
horizontal rows of regular pores. Consequently, the wall tissue between neighbouring
332
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
horizontal rows of pores, forms a continuous circular bar. A small tongue-like plate
arises from the bar under each pore, but it was not possible to discover whether these
tongues are separate from each other, or connected to form an annular ring. The
radial septa have scattered regular pores, while the tabulae consist of a thin porous
net.
DIMENSIONS
Cup:
Height
Diameter .....
Intervallum ....
Interseptum (outer wall)
(inner wall)
Intervallum coefficient
Parietal coefficient
Outer wall :
8 pores (4 + 4) per interseptum
Diameter of pores
Thickness of linteaux .
Thickness of wall
Inner wall :
No. of rows of pores per interseptum
Diameter of pores
Vertical partitions
Horizontal partitions .
Thickness of wall
Septa :
No. of rows of pores per septa
Diameter of pores
Vertical partitions
Horizontal partitions .
Tabulae :
No. of rows of pores per interseptum
Diameter of pores
Partitions .
(mm.)
9
9
i'5i
0-83
0-56
0-25
0-07
O'Oy
O'll
2
0-07
O-II
0-07
0-26
6
0-03
o-ii
• 11-0-26
12
0-03
O-II
DISCUSSION. The horizontal alignment of the inner wall pores is an important
feature, that is a precursor to the formation of annular structures. This is evident in
this case, for the horizontal bars are already slightly modified by supplementary
plates. However, as the stage of a single pore per interseptum has not been reached,
this species should be retained in Erismacoscinus and not placed in a genus of the
Salairocyathidae. The star-shaped form of the external wall is similar to that of E.
cancellatus (Bornemann), but the two species have no other features in common.
Erismacoscinus retifer (R. & W. R. Bedford)
(PL 10, figs. 2 and 3)
1934 Coscinocyathus retifer R. & W. R. Bedford : 3, pi. 3, fig. 14.
1964 Erismacoscinus retifer (R. & W. R. Bedford) Debrenne : 167.
HOLOTYPE. B.M. (N.H.) S 4159.
LOWER CAMBRIAN ARCH AEOC Y ATM A FROM AUSTRALIA 333
DESCRIPTION. The species is based on a single, small, incomplete specimen, but it
is sufficiently well-preserved to enable one to see the details of each skeletal plate.
The thin outer wall is perforated by 4-5 vertical rows of pores to each interseptum,
these pores also form horizontal rows. The septa and the only visible tabula are of
the same thickness as the outer wall, and have their round pores arranged in
quincunx; the skeletal tissue between them is wider than the pores and projects
slightly.
The inner wall is thicker than the other plates. The pores are elliptical, with the
larger diameter horizontal and a tongue arising under each pore. One vertical row of
pores is only just separated from the next and, at times, pores from neighbouring
rows may join, giving a single pore at that point of the interseptum. At certain
places, the pores are aligned horizontally and this involves two sporadic modifications.
Firstly, the coalescence of two neighbouring pores, which predicts the uniserially-
pored inner wall. Secondly, the development of small tongue-like plates under the
pores, forecasting the formation of annular plates.
DIMENSIONS
(mm.)
Cup:
Height (pars) ...... 9
Diameter ...... 8
Intervallum . . . . . . 1-96
Intervallum coefficient . . . . 0-5
Parietal coefficient . . . . . 3-5
Outer wall:
No. of pore rows per interseptum . . . 4-5
Diameter of pores . . . . . 0-07
Vertical partitions . . . . . o-io
Horizontal partitions . . . . . o-io
Thickness . . . . . . . 0-13
Inner wall :
Diameter of pores ..... 0-28-0-18
Vertical partitions . . . . o-io
Horizontal partitions . . . . . 0-15
Length of plates . . . . . . 0-20
Septa :
No. of pores per loculus .... 9-10
Diameter of pores . . . . . o • 1 1
Distance between horizontal rows . . o • 1 8
Distance between vertical rows . . . 0.22
Tabulae :
No. of pores per interseptum . . . 6
Diameter . . . . . . . 0-04
Partitions . . . . . . . 0-05
Thickness . . . . . . . 0-05
DISCUSSION. This species continues to follow the evolution already outlined under
E. peter si and heralds the annular forms of Salairocyathidae.
334 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
Family POLYCOSCINIDAE Debrenne, 1964
DIAGNOSIS. Cup with porous septa and tabulae. Double, porous outer wall.
Simple inner wall.
COMPOSITION OF THE FAMILY. Polycoscinus R. & J. Bedford 1937, Tomocyathus
Rozanov 1960
Genus TOMOCYATHUS Rozanov, 1960
1960 Tomocyathus Rozanov : 664, figs, i d, e.
1964 Tomocyathus Rozanov; Debrenne : 115
1964 Tomocyathus Rozanov; Repina et al. : 231.
1965 Tomocyathus Rozanov; Hill : 107
TYPE SPECIES. Tomocyathus operosus Rozanov 1960, by original designation.
DIAGNOSIS. Outer wall coarsely porous, covered by a secondary micro-porous
sheath. Straight septa and convex tabulae, both with numerous fine pores. Vesi-
cular tissue. Inner wall with simple pores, and stellate in transverse section.
DISCUSSION. The fluting of the walls is often considered a character of sub-
generic value (Krasnopeeva 1955; Debrenne 1964). Recently, Repina et al (1964 :
231) described Tomocyathus without any mention of the stellate inner wall. Some
species that they included in this genus, compositus (Zhuravleva) and shoriensis
Rozanov, have little to no vesicular tissue, rather flat tabulae and the inner wall is not
fluted but possesses spines of varying size and shape.
According to criteria used in distinguishing other subgenera e.g. Ajacicyathus
(Urcyathus) Vologdin, one must create a new subgenus for the smooth forms of
Tomocyathus.
COMPOSITION OF THE GENUS. Tomocyathus (Tomocyathus) operosus Rozanov 1960,
T. (Tomocyathus} michniaki Rozanov 1966; for subgenus Tomocyathus (Erugato-
cyathus} see below.
Subgenus ERUGATOCYATHUS nov.
DERIVATION, erugatus — unfolded.
TYPE SPECIES. Coscinocyathus papillatus R. & W. R. Bedford, designated here.
DIAGNOSIS. Tomocyathus with non-folded inner wall.
COMPOSITION OF THE SUBGENUS. Tomocyathus (Erugatocyathus} echinus (Debrenne
1964), T. (E.) compositus Zhuravleva 1955, T. (E.) gini Missarzhevsky & Rozanov
1962, T. (E.) kundatus Rozanov 1966, T. (E.) shoriensis Rozanov 1964, T. (E.) papilla-
tus (R. & W. R. Bedford 1934).
Tomocyathus (Erugatocyathus) papillatus (R. & W. R. Bedford)
(PL n, fig. 2, Text-fig. 9)
1934 Coscinocyathus papillatus R. & W. R. Bedford : 3, pi. 3, fig. 12.
1964 Erismacoscinus papillatus (R. & W. R. Bedford) Debrenne : 167
HOLOTYPE. B.M. (N.H.) S 4153.
PARATYPE, B.M. (N.H.) S 4154.
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
335
DESCRIPTION. Fragments of cylindrico-conical fossils. Septa with remote round
pores. Tabulae reticular. Loculi rectangular and irregular in size. The circular
pores at the base of the coarsely-porous outer wall are covered by a microporous
sheath, in which each group of micropores consists of a central pore surrounded by
five others.
A skeletal tongue covers each simple pore of the inner wall, it arises under the pore,
curves upwards and rejoins the wall above. As the rows of pores alternate, each pore
is surrounded by 4 knobs, corresponding to the beginning of a papilla. This feature is
well shown in the weathered specimen S 4154 (see Text-fig. 9).
FIG. 9. Tomocyathus (Erugatocyathus) papillatus (R. & W. R. Bedford)
DIMENSIONS
Cup:
Height
Diameter
Intervallum coefficient
Parietal coefficient
Intervallum
Interseptum
S 4153 Upper S 4153 Lower
(mm.) (mm.) (mm.)
13
1-89
3-8
0-40
o-34
18
10-5
0-41
2-5-4
336
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
DIMENSIONS — continued
Outer wall:
No. of rows of pores per interseptum
Diameter
Vertical skeletal partitions
Horizontal skeletal partitions
Thickness
Inner wall:
No. of pore rows per interseptum
Diameter
Vertical skeletal partitions
Horizontal skeletal partitions
Thickness
Septa :
No. of pore rows per inter vallum
Diameter
Vertical skeletal partitions
Horizontal skeletal partitions
Thickness
Tabulae
No. of pore rows per interseptum
Diameter
Vertical skeletal partitions
Thickness
S 4153 Upper
(mm.)
84153 Lower
(mm.)
S4I54
(mm.)
2
2
0-15
0-15-0- 18
0-15-0-03
0-15-0-03
0-18
0-15
0-15-0-03
0-15
0-15
O-II
2
2
2
O-22
0-20
0-20
0-15
0-15
o-io + pap. = o-n 0-22
10 8
0-05 0-07
o-io o-ii
0-07 0-07
5-6 4-5
0-05 0-05
0-07 0-05
0-05 0-05
0-07
o-ii
O-II
0-07
4
0-07
0-07
0-05
DISCUSSION. These peculiar curved tongues are not known in any other species
of the Regularia.
Family SALAIROCYATHIDAE Zhuravleva, 1955
DIAGNOSIS. Cups with intervallum crossed by porous septa and tabulae. Simple
outer wall, but annular inner wall.
COMPOSITION OF THE FAMILY. Salairocyathus (Salairocyathus) Vologdin 1940,
Salairocyathus (Polystillicidocyathus} Debrenne 1959.
Genus SALAIROCYATHUS Vologdin, 1940
i94oa Salairocyathus Vologdin : 89
TYPE SPECIES. Salairocyathus zenkovae Vologdin 1940 : 89, pi. 26, fig. 6.
DIAGNOSIS. Simple pores in outer wall and also in septa and tabulae. Annuli on
inner wall, v-shaped in section and open towards the top.
COMPOSITION OF THE GENUS. Salairocyathus (Salairocyathus} zenkovae Vologdin
1940, 5. (S.) pospelovi Zhuravleva 1960, ? 5. (S.) annulatus (R. & W. R. Bedford
1934). Salairocyathus (Polystillicidocyathus} erbosimilis Debrenne 1959.
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 337
? Salairocyathus (Salairocyathus) annulatus (R. & W. R. Bedford)
(PI. ii, fig. 4, Text-fig. 10)
1934 Coscinocyathus annulatus R. & W. R. Bedford : 4, pi. 3, fig. 17.
HOLOTYPE. S 4163 in British Museum (Natural History).
DESCRIPTION. A unique, very fragmentary specimen, that has part of the inner
wall, as seen from the central cavity, well preserved. Porous outer wall, but not
FIG. 10. Salairocyathus (Salairocyathus} annulatus (R. & W. R. Bedford)
possible to distinguish their arrangement. Septa and tabulae frame somewhat
parallelepipedic loculi. The rectangular openings of the inner wall are bounded by
the inner edges of the septa and a vertical series of horizontal bars, which are
annular in form. The ft component of the latter is thick and horizontal, while the a
component is much thinner and curves backwards up to the middle of the following
horizontal row of openings, which are thus partly closed, (see Text-fig. 10).
338 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
DIMENSIONS
(mm.)
Cup:
Height 27
Diameter 1 1
Intervallum ?
Interseptum o • 22-0 • 30
Intertabulum about 1-35
Outer wall:
No. of rows of pores per interseptum probably 4
Thickness 0-13
Inner wall:
No. of rows of pores per interseptum i
Diameter of pores (lengthened horizontally) 0-25-0 -15
Skeletal partitions (constant thickness) o • 06
Septum :
Diameter of pores 0-06
Skeletal partitions 0-13
Thickness 0-13
Tabulae :
Diameter of pores (radially lengthened) 0-13-0-06
Skeletal partitions o • 06
Thickness 0-15
DISCUSSION. This annular form of inner wall was previously unknown. The
specimen is described as Salairocyathus with reservation.
Family ALATAUGYATHIDAE Zhuravleva, 1955
DIAGNOSIS. Cups with simply porous septa and tabulae. Inner wall with simple,
or S-shaped pores. Two sub-families are distinguished by the shape of the pores in
the outer wall.
Subfamily TUMULOCOSCININAE Zhuravleva, 1960
DIAGNOSIS. Outer wall with simple tumuli.
COMPOSITION OF THE SUBFAMILY. Tumulocoscinus Zhuravleva 1960, Ethmoco-
scinus Simon 1939, Asterotumulus Rozanov 1964.
Subfamily ALATAUGYATHINAE Zhuravleva, 1960
DIAGNOSIS. Outer wall with knobbly tumular pores.
COMPOSITION OF THE SUBFAMILY. Alataucyathus (Alataucyathus) Zhuravleva
1955, Alataucyathus (Anaptyctocyathus) subgen. nov.
Subfamily TUMULICOSCININAE Zhuravleva, 1960
Genus ETHMOCOSCINUS Simon 1939
1939 Ethmocoscinus Simon : 28
TYPE SPECIES. Coscinocyathus papillipora R. & W. R. Bedford, by original desig-
nation (Simon 1939 : 28).
LOWER CAMBRIAN ARCH AEOCYATH A FROM AUSTRALIA 339
DIAGNOSIS. Cylindro-conical cups. Outer wall with simple tumuli, one per
interseptum. Inner wall with a single row of S-shaped pore-tubes per interseptum.
Septa have sparse simple pores. Tabulae with polygonal, somewhat irregular
pores.
COMPOSITION OF THE GENUS. A single species E. papillipora (R. & W. R. Bedford).
Ethmocoscinus papillipora (R. & W. R. Bedford)
(PL 3, fig- 3)
1934 Coscinocyathus papillipora R. & W. R. Bedford : 18.
1939 Ethmocoscinus papillipora (R. & W. R. Bedford) Simon : 28.
HOLOTYPE. B.M. (N.H.) S 4164.
DESCRIPTION. The holotype, the only known specimen, is partly weathered-out
and shows the structure of the two walls. The straight septa are perforated by round
regular pores, which although few, are arranged in quincunx. A single tabula is
visible (it is therefore impossible to state the frequency) and consists of a net of regular
pores. The papillae of the outer wall are situated between the septa in vertical rows,
which are just separate from one interseptum to the next. The papillae of the inner
wall occupy a much larger area and also appear to be in alternate vertical rows.
DIMENSIONS
Cup: (mm.)
Height (pars) 20
Diameter 7 • 5
Intervallum i • 2
Interseptum from 0-41-0-45
Number of septa 26
Parietal coefficient 3 • 4
Intervallum coefficient 0-2
Outer wall:
No. of pore rows per intersept i
No. of pore rows before formation of septa 2
Diameter of papilla o • 34-0 • 2 2
Thickness of papilla o • 07
Height of papilla o • 18
Horizontal partitions 0-15-0-18
Vertical partitions o • 1 1
Inner wall:
No. of rows of pores per interseptum I
Diameter of pores 0-37
Partitions o • 1 1
Septa :
No. of rows of pores per intervallum 4
Diameter of pores o • 1 1
Partitions o • 1 1
Thickness o • 06
Tabula :
No. of rows of pores per interseptum 6
Diameter of pores o • 07
Partitions o • 03
Thickness o • 03
340 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
Subfamily ALATAUCYATHINAE Zhuravleva, 1960
Genus ALA TA UCYA THUS Zhuravleva, 1955
1955 Alataucyathus Zhuravleva : 626, figs. la, 2d.
TYPE SPECIES. By original designation Alataucyathus jaroschevitschi Zhuravleva
1955 : 626.
DIAGNOSIS. Intervallum filled with simple septa and tabulae. Outer wall covered
with small multi-perforate knobs. Inner wall simple with vertical folds between
neighbouring septa.
DISCUSSION. As mentioned in the discussion on Tomocyathus, it is considered that
species having non-fluted inner walls, should be placed in a separate sub-genus.
Subgenus ALATAUCYATHUS (ANAPTYCTOCYATHUS) nov.
DERIVATION, anaptyctos = unfolded.
TYPE SPECIES. Coscinocyathus cribripora R. & W. R. Bedford designated here.
COMPOSITION OF THE GENUS. A. (A.) cribripora (R. & W. R. Bedford, 1934), A . (A .)
minimiporus (R. & J. Bedford 1937) — included with some doubt as the structure of
the walls is not certain, A. (A.) excellenthis (Rozanov 1964), A. (A.) verschkhovska-
jae (Zhuravleva 1961), and A . (A .}flabellus nov.
Alataucyathus (Anaptyctocyathus] cribripora (R. & W. R. Bedford)
(PL n, figs, i and 3)
1934 Coscinocyathus cribripora R. & W. R. Bedford : 3, pi. 3, fig. 15.
HOLOTYPE. B.M. (N.H.) S 4160.
DESCRIPTION. Cylindrical cup. Intervallum with straight radial septa, in which
the hexagonal pores occupy a much larger surface in proportion to the skeletal tissue,
and irregularly distributed flat tabulae, that are finely perforated by very narrow
pores.
The inner wall is simple, with two rows of pores per interseptum, which are only
weakly separated, in fact, towards the upper part of the cup, two neighbouring pores
may sometimes coalesce. The pores of the outer wall are covered by knobs, that are
perforated by a central pore and a surrounding circle of six others.
DIMENSIONS
(mm.)
Cup:
Height (pars) 36
Upper diameter 12
Lower diameter 6
Intervallum coefficient 0-28
Interseptum (Upper diameter) °'74
(Lower diameter) o • 56
Intertabula 2-5
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA 341
DIMENSIONS — continued
(mm.)
Outer wall:
No. of rows of main pores per interseptum 2
(each pore covered by a sheath with 7 micropores)
Main pore diameter 0-27
skeletal partitions o • 27-0 • 30
Micropore diameter 0-07
skeletal partitions o • 03
Inner wall :
No. of pore rows per interseptum 2
(these sometimes coalesce)
Diameter of pores o • 30
Skeletal partitions o • 07
Septa:
No. of hexagonal pores 4-5
Diameter o • 1 5-0 • 1 8
Vertical partitions o • 18
Horizontal partitions 0-18
Tabulae :
No. of pores 10-4
Diameter o • 1 1 -o • 06
Partitions o • 07
DISCUSSION. A. (A.) cribripora differs from the two Russian species by having
lower tumuli, coarse micropores and a thicker inner wall.
Alataucyathus (Anaptyctocyathus) flabellus nov.
(PI. 10, fig. i)
HOLOTYPE. B.M. (N.H.) S 4161.
DESCRIPTION. Bowl-shaped cup with narrow intervallum. Regularly porous
septa in alternate vertical rows. The tabulae almost join, their pores are much
thinner than those of the septa. The inner wall is only known from the intervallum
side and therefore, it is not possible to discover whether there are spines on the central
cavity face. Three vertical rows of pores can be distinguished per interseptum. The
outer wall has 2-3 pores, each covered by a small microporous tumulus with about ten
angular pores in each.
DIMENSIONS
(mm.)
Cup:
Height (pars) 20
Span of visible arc 78
Intervallum 2
Loculi 1/2/3
Outer wall:
No. of rows of pores per interseptum from 2-3
Diameter of main pores 0-40-0-27
342 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
DIMENSIONS — continued
(mm.)
Vertical skeletal partition o • 16
Horizontal skeletal partition o • 1 1
Diameter of micropores o • 06
Skeletal partition between micropores 0-03
Inner wall:
Diameter of pores 0-18
Skeletal partitions o • 22
Thickness 0-20
Septa :
Diameter of pores o • 20
Vertical partitions o • 30
Horizontal partitions 0-20
Thickness o-n
Tabulae :
No. of rows of pores per interseptum 5
Diameter of pores 0-06
Skeletal partitions o • 06
Thickness o • 1 1
DISCUSSION. Differs from A. (A.) cribripora by its bowl-shaped cup, the specific
coefficients and the different arrangement of the outer wall pores.
Class IRREGULARIA Vologdin, 1937
Order ARCHAEOCYATHA Okulitch, 1935
Family DICTYOGYATHIDAE Taylor, 1910
DIAGNOSIS. Cups with intervallum containing disorientated short rods and dis-
sepiments. Two simply porous walls.
COMPOSITION OF THE FAMILY. Dictyocyathus Bornemann 1891, Spinosocyathus
Zhuravleva 1960, Pinacocyathus R. & W. R. Bedford 1934, Agastrocyathus Debrenne
1964, Archaeopharetra R. & W. R. Bedford 1936, Chouberticyathus Debrenne 1964.
Genus PINACOCYATHUS R. & W. R. Bedford, 1934
1934 Pinacocyathus R. & W. R. Bedford : 4, fig. 21.
1964 Pinacocyathus R. & W. R. Bedford; Debrenne : 200
1965 Pinacocyathus R. & W. R. Bedford; Hill : 117, fig. 22, ya and b.
TYPE SPECIES. By monotypy Pinacocyathus spicularis R. & W. R. Bedford
1934 : 4.
DIAGNOSIS. Two-walled cup with intervallar rods. The outer wall is formed by a
scaffolding of vertical pillars connected by horizontal, or slightly oblique rods. The
inner wall is, probably, a regular net. Oblique, or radial horizontal rods in the
intervallum.
COMPOSITION OF THE GENUS. A single species; P. spicularis.
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
343
Pinacocyathus spicularis R. & W. R. Bedford
(PI. 12, figs, i, 2, Text-fig, n)
1934 Pinacocyathus spicularis R. & W. R. Bedford : 4, fig. 21.
HOLOTYPE. By monotypy B.M. (N.H.) S 4169.
DESCRIPTION. Conical tube. The outer wall consists of an open framework,
which is mainly composed of vertical columns arranged around the cup at almost
regular intervals. This framework is completed by horizontal, or oblique rods,
FIG. ii. Pinacocyathus spicularis R. & W. R. Bedford
which connect the columns, with a few spines occurring at the junctions. On the
lower part, the net is more compact but hidden by poor preservation, that has
swollen the structures. Only one vertical row of the inner wall can be seen and this
seems to be composed of a regular mesh. However, it was not possible to determine
the arrangement of the intervallar rods. They seem to arise from the skeletal junc-
tions of the outer wall framework and continue radially, or obliquely towards the
inner wall. The framework is probably thicker in its lower parts.
GEOL. 17, 7 27
344 LOWER CAMBRIAN ARCH AEOC Y ATM A FROM AUSTRALIA
DIMENSIONS
(mm.)
Cup:
Height (pars) . . . . . . 15
Diameter . . . . . . . 5 • 75
Intervallum . . . . . . 1-42
Central cavity . . . . . . 0-6
Outer wall :
Diameter of pores . . . . . o • 55
Distance between vertical rods . . . o • 67-1 • o
Distance between horizontal rods . . . i • 35-1 • 70
Thickness of vertical rods . . . . °'33
Thickness of horizontal rods . . . 0-33
Thickness of oblique rods . . . . 0-25
Inner wall :
Diameter of pores . . . . . °'33
Vertical partitions . . . . . 0-20
Horizontal partitions • • • • • 0-20
Thickness . . . . . . . 0-20
Intervallar rods :
Thickness . . . . . . . 0-27
DISCUSSION. Only one specimen of this curious form is known. It is included in
the Dictyocyathidae because of its scaffolding of intervallar rods, but no other
recognized genus has such a loosely-spaced framework.
Family FLINDERSICYATHIDAE R. & J. Bedford, 1939
DIAGNOSIS. Two-walled cups, simply porous, with a single series of large pores in
the inner wall. A scaffolding of rods fills the intervallum, consisting of taeniae, that
cross the intervallum, joining the septa, and undulate in a radial plane, together with
synapticulae, which connect the crests of neighbouring taeniae. Little to no vesicular
tissue is present. Apex of Dictyocyathus-type.
COMPOSITION OF THE FAMILY. Spirocyathella Vologdin 1939, Flindersicyathus
(Flindersicyathus} R. & J. Bedford 1937, Flindersicyathus (Pycnoidocyathus} (Taylor
1910), ? Spirillicyathus R. & J. Bedford, 1937, Copleicyathus R. & J. Bedford 1937.
Genus FLINDERSICYATHUS R. & J. Bedford, 1937
1910 Pycnoidocyathus Taylor : 131.
1937 Flindersicyathus R. & J. Bedford : 28.
1939 Flindersicyathus R. & J. Bedford; Simon : 30.
1960 Archaeocyathus Billings; Zhuravleva : 296 (pars).
1965 Flindersicyathus R. & J. Bedford; Hill : 123.
TYPE SPECIES. Flindersicyathus decipiens R. & J. Bedford, by subsequent
designation R. & J. Bedford (i939(May) : 78), which has priority over Spirocyathus
irregularis Taylor chosen by Simon (1939 (Dec.) : 30), see Hill (1965 : 123). Holo-
type material No. 86670 in Princeton University, U.S.A.
DIAGNOSIS. Hill (1965 : 123) writes: "solitary cups, with simply porous outer
wall; inner wall with a single series of large rounded pores per intertaenial loculus,
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
345
each pore bounded by the taeniae and the synapticulae joining them, and each with a
louvre-like plate projecting upwards and inwards to the central cavity from its low
synapticula". [In fact, each pore is a short tube with the lower part projecting into
the central cavity, rather than a simple pore with a projecting plate.] Intervallum
with taeniae, sparsely porous near the inner wall, coarsely and copiously porous else-
where; the taeniae are waved in the radial plane, and the waves have angulated
crests and troughs, the crests and the trough-lines curving upwards and outwards
FIG. 12. Archaeocyathus atlanticus Billings X 4
from the inner wall ; the crests of neighbouring taeniae are opposed and connected by
synapticulae. Rare dissepiments may occur. No tabulae ".
DISCUSSION. R. & J. Bedford (1937 : 28) and Hill (1965 : 123 and 128) point out
that Flindersicyathus R. & J. Bedford and Pycnoidocyathus Taylor have a very similar
structure. The latter differs only in having much stronger transverse annulations of
the intervallum, the inner wall of both remaining cylindrical. The two genera may
be synonymous, but I have not seen examples of the type species of Flindersicyathus,
or the type material of Pycnoidocyathus and therefore cannot settle the question. My
tentative suggestion is that Pycnoidocyathus should be considered a subgenus of
Flindersicyathus, due to its different outer shape, although possibly having a similar
structure. This situation is comparable to the distinction between Ajacicyathus R. &
346 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
J. Bedford and Orbicyathus Vologdin, which are considered to be subgenera distin-
guished by their external shape.
Zhuravleva (1960 : 296) placed Flindersicyathus in synonymy with Archaeocyathus
Billings 1861. Debrenne (1964 : 117) doubtfully placed Flindersicyathus in the family
Archaeocyathidae (as did Hill 1965 : 123), but had maintained its independence.
However, having seen the holotype of Archaeocyathus atlanticus Billings (1861, fig. 10).
I can state that the taeniae are thickened by several skeletal layers, have few and
irregular pores (Text-fig. 12), are goffered longitudinally and transversely, and are
occasionally connected to the opposite crests, more or less forming elongated pores.
The curved hexagonal tubular mesh of Flindersicyathus was not recognized. Further
characters which separate the two genera are the abundant vesicular tissue, the pore
canals of the inner wall and the irregular net of the outer wall of Archaeocyathus.
COMPOSITION OF THE GENUS. Flindersicyathus (Flindersicyathus) : F. decipiens R.
& J. Bedford, 1937, F. circliporus R. & J. Bedford, 1937, F. contractus Hill 1965, F.
graphicus (R. & W. R. Bedford 1934), F. irregularis (Taylor 1910), F. latiloculatus
Hill 1965, F. major (R. & W. R. Bedford 1934), F. multifidus (R. & W. R. Bedford
1936), ? F. macdonnelli R. & J. Bedford 1937, F. rete (R. & W. R. Bedford 1936),
F. simplex (Taylor 1910), F. speciosus (R. & W. R. Bedford 1934), F. tabulatus R. &
J. Bedford 1937.
Flindersicyathus (Pycnoidocyathus) : F. synapticulosus (Taylor 1910), F. maximipora
(R. & W. R. Bedford 1936), F. parvulus (R. & W. R. Bedford 1936), F. ptychophragma
(Taylor 1910), F. vicinisepta (R. & W. R. Bedford 1936).
\
Flindersicyathus (Flindersicyathus) graphicus (R. & W. R. Bedford)
(PI. 12, figs. 3-5, Text-fig. 13)
1934 Protopharetra graphica R. & W. R. Bedford : 4, pi. 4, fig. 22.
1939 Dictyocyathus graphicus (R. & W. R. Bedford) R. & J. Bedford : 73.
1964 Metaldetes graphica (R. & W. R. Bedford) Debrenne : 220.
HOLOTYPE. B.M. (N.H.). S 4170.
PARATYPES. B.M. (N.H.) S 4171-4173.
OTHER MATERIAL. B.M. (N.H.) S 4761, S 7629.
DESCRIPTION. Slender conical cups, undulating exterior form, with the bulges of
the inner wall following those of the outer wall, so that the intervallum remains a
constant width. Thin outer wall, perforated by hexagonal to ellipsoidal pores, re-
calling the outer wall of the Regularia. There is no specific inner wall, but, instead,
the innermost face is formed by the inner edges of the taeniae and connecting bars,
which enclose apertures that are, in effect, " wall pores ".
The structure of the intervallum can, perhaps be considered as an assemblage of
septa of the Volvacyathus-type, with regular, somewhat rounded, hexagonal pores set
in a radial plane, with the line of pores curving upwards and outwards from the inner
to the outer wall. These pseudo-septa (flat taeniae) are joined to one another by
horizontal synapticulae and join the skeleton at each pore angle. No vesicular tissue.
Early stages of the species unknown.
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
347
FIG. 13. Flindersicyathus (Flindersicyathus} graphicus (R. & W. R. Bedford)
DIMENSIONS
Cup:
Height
Diameter
Intervallum
Intervallum coefficient
Bars:
Radial distance
Tangential distance
Vertical distance
Thickness
Outer wall :
No. of pores per
intertaeniae
Diameter of pores
Partitions
Thickness
Inner wall:
No. of pores per
intertaeniae
Diameter of pores
Partitions
Thickness
S 4170
(mm.)
35
12
S 4171
(mm.)
45
S 4172
(mm.)
15
15
sup.
(mm.)
7
inf.
(mm.)
4
S 7629
(mm.)
45
20
S 4761
(mm.)
30
18
1-9
1-7
2
i -5
i -13
i -5
1 '5
0-25
0-17
0-18
o-3
0-6
o- 1
O' I
0-26
0-18
o-34
0-15
0-15
o-37
0-22
0-26
0-26
0-41
O-22
0-30
0-30
0-22
o-37
0-26
o-37
0-36
O-22
0-07
0-07
0-07
0-07
0-07
0-07
O-O7
0-11-0-09
0-03
o- ii
I
0-26
0-13
0-18
3
0-07
0-03
0-07
i
O-22
0-07
O-II
o-37
0-15
0-05
I
0-18
0-07
O-II
3
o-ii
0-05
0-07
0-18
o-ii
3
o-ii
0-07
0-07
0-18
o- ii
0-07
348 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
DISCUSSION. The taeniae (pseudosepta) of graphicus are not waved in the radial
plane, as has been described in F. decipiens (R. & J. Bedford) probably because of the
narrowness of the intervallum and the density of the taeniae and synapticulae. The
inner wall does not have a plate on the lower part of the pore. Thus graphicus is much
simpler than other species of Flindersicyathus, but the presence of well defined radial
plates (flat taeniae) indicates a more complex stage than that of having sparse rods or
plaquettes.
The species was first described as Protopharetra, subsequently as Dictyocyathus and
then, in 1964, removed to Metaldetes because of the pseudosepta. However, it differs
from Metaldetes in having a very regular outer wall, numerous synapticulae and the
absence of connections between these. The scaffolding of the intervallar mesh is very
like that of Tabellaecyathus Fonin but the walls are simpler, neither tabello-reticulate
nor tabello-porous.
Flindersicyathus (Flindersicyathus} irregularis (Taylor)
(PI. 14, fig. 4)
1910 Spirocyathus irregularis Taylor : 148, pi. 16, figs. 93-94.
1936 Spirocyathus irregularis Taylor; R. & W. R. Bedford : 14, pi. 13, fig. 64.
1937 Flindersicyathus irregularis (Taylor) R. & J. Bedford : 28.
J937 Spirocyathus atlanticus Billings; Ting : 368-369, pi. 13, fig. 14.
1961 Archaeocyathus irregularis (Taylor) F. & M. Debrenne : 702, pi. 20, fig. 5.
MATERIAL. B.M. (N.H.) S 7625, S 4763.
DESCRIPTION. Two-walled cup with numerous thin radial taeniae, probably quite
porous [a deduction made from Taylor's comparison with F. rete in his original
description of F. irregularis and not from observation, as no specimen showing a
longitudinal section was available], wavy and connected by synapticulae which
form a network of more or less regular polygonal apertures, particularly in the outer
part. Vesicular tissue sparse. Skeletal tissue is sometimes thickened by secondary
layers and occurs mainly near the inner wall (see PI. 14, fig. 4; also F. & M. Debrenne,
1961, pi. 20, fig. 5). Thin outer wall, with numerous small pores; inner wall with one
vertical row of pores per intertaenial space.
DIMENSIONS
Taylor Ting S 7625 S 4763
(mm.) (mm.) (mm.) (mm.)
Cup:
Height (pars) 50 17 16 25
Diameter 14 16-5 18 20
Intervallum coefficient 0-5 0-8 0-8 0-9
Intertaenial space 0-5 0-4 ° ' 45 ° ' 5
Intersynapticular space
Horizontal . . . . 0-70 0-6
Vertical . . . . irregular
Taeniae thickness o • i 0-12 0-2 0-2
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 349
DIMENSIONS — continued
Taylor Ting S 7625 S 4763
Outer wall: (mm-) (mmO (mm-) (mm-)
Diameter of pores .. 0-2-0-4
Partitions .. 0-12
Thickness .. 0-12 0-13 0-15
Inner wall :
Diameter of pores . . 0-5 0-35 o • 33
Partitions .. 0-3 0-27 0-3
Thickness . . i i i
DISCUSSION. The taeniae are thinner and more numerous than in other species of
Flindersicyathus, and the thickening of the inner wall recalls some transverse sections
of Copleicyathus confertus R. & J. Bedford (Hill 1965: pi. 10, figs. 4a-b), but at the
moment, it is not possible to take the comparison any further.
Flindersicyathus (Flindersicyathus) major (R. & W. R. Bedford)
(PL 14, fig. 2)
1934 Spirocyathus major R. & W. R. Bedford : 5, pi. 4, fig. 24.
HOLOTYPE. B.M. (N.H.) S 4174.
DESCRIPTION. Only one broken specimen known. The outer wall is obscured by
silicification. The inner wall has round, shaft-like pore tubes, that are limited by
neighbouring taeniae and successive vertical synapticulae ; the lower part of each tube
is inclined downwards into the intervallum and projects slightly into the central
cavity, whereas the upper part is flattened. The rows of pore-tubes alternate. The pores
are more crowded at the base of the cup, because the taeniae are closer together there.
The intervallum is crossed by waved taeniae, with the crests and troughs of neigh-
bouring taeniae opposite to one another. Abundant synapticulae join the taeniae,
particularly in the central and outer parts of the intervallum. The ratio diameter of
taenial pores : distance apart of the synapticulae is close to unity, so that the inter-
vallum has the appearance of a tubular mesh with tubes curving upwards and
outwards.
DIMENSIONS
Cup: (mm.)
Height (pars) ...... 40
Diameter ....... about 20
Intervallum ...... 10
Interseptum . . . . . . 1-5
Outer wall :
Thickness . . . . . . 0-20
Inner wall :
Diameter of pores : horizontal . . . 1-70
vertical . . . i-I5
Horizontal partitions . . . . i'i5
Lateral partitions . . . . . 0-33
Tubes :
Irregular diameter ..... about o • 67
Large skeletal plates
Thickness . . . . . . 0-27
350 LOWER CAMBRIAN ARCH AEOC Y ATM A FROM AUSTRALIA
DISCUSSION. The septa are at wider intervals than in other species of Flinder-
sicyathus.
Flindersicyathus (Flindersicyathus) speciosus (R. & W. R. Bedford)
(PL 14, figs, i, 3)
1934 Spirocyathus speciosus R. & W. R. Bedford : 5, pi. 4, fig. 25.
HOLOTYPE. B.M. (N.H.) S 4175.
DESCRIPTION. Only a single broken specimen known. The taeniae are thin, radial,
slightly waved and connected by sparse synapticulae, which are chiefly found in the
outer part of the cup. The diameter of the taenial pores is less than the width of the
skeletal tissue between them.
Outer wall with 4 to 5 alternating vertical rows of round pores per interseptum.
The inner wall has a single tube per intertaenial space, the lower part of which projects
into the central cavity, giving the tube a crescentic section. The rows alternate.
DIMENSIONS
Cup: (mm.)
Height (pars) ...... 30
Upper diameter ..... about 13 -5
Lower diameter ..... 1 1
Intervallum . . . . . . 4-5
Intervallum coefficient .... i
Intertaenial space ; lower end of cup . . 0-6
upper part of cup . . o • 75
Outer wall :
Diameter of pores . . . . . 0-15
Vertical skeletal partitions . . . . 0-07
Horizontal skeletal partitions . . . 0-18
Inner wall :
Diameter of pores ; horizontal . . . 0-9
vertical . . . 0-4
Vertical skeletal partitions . . . . 0-13
Crescentic plates . . . . . 0-33
DISCUSSION. The taeniae are less waved and not so distinct as those in Flinder-
sicyathus major. The shaft-like tubes of the latter are not present, while the inner
wall pores of speciosus are crescentic and very close together. Synapticulae are less
numerous than in other Flindersicyathus species.
Subgenus FLINDERSICYATHUS (PYCNOIDOCYATHUS) (Taylor), 1910
1910 Pycnoidocyathus Taylor : 132.
1939 Pycnoidocyathus Taylor; R. & J. Bedford : 78.
1965 Pycnoidocyathus Taylor; Hill : 128
TYPE SPECIES. P. synapticulosus Taylor (1910 : 132) by subsequent designation
R. & J. Bedford, (1939 : 78) ; type material in the Univ. of Adelaide.
DISCUSSION. Although Hill (1965 : 128) has provisionally treated this as a genus
separate from Flindersicyathus, I regard it as a subgenus (see p. 345). It is very like
Flindersicyathus, but with much stronger expansions and contractions of the inter-
LOWER CAMBRIAN ARCH AEOCY ATM A FROM AUSTRALIA 351
vallum which do not affect the inner wall. The problem can only be settled by exami-
nation of the type material, in order to discover whether dissepiments occur and the
exact nature of the inner wall pores.
Flindersicyathus (Pycnoidocyathus) synapticulosus (Taylor)
(PI. 15, fig- i)
1910 Pycnoidocyathus synpaticulosus Taylor : 132, pi. 12, fig. 69.
1936 Pycnoidocyathus synapticulosus Taylor; R. & W. R. Bedford : 15, pi. 15, fig. 69.
HOLOTYPE. Probably at Adelaide University.
MATERIAL. B.M. (N.H.) S 208, S 4825.
DESCRIPTION. Large cup with annular bulges. On the specimens examined here,
the outer part of each bulge is made of successive flanges. Outer wall with somewhat
irregular polygonal fine pores. Inner wall with a vertical series of short tubes per
intertaenial space, the tubes lead upwards into the central cavity. Taeniae are
radial, nearly flat and close together, particularly near the synapticulae, so that
transverse sections appear to be composed of irregular polygonal cells. Abundant
horizontal and vertical synapticulae ; generally arranged in quincunx from one inter-
sept to the next. It was not possible, to examine the size and arrangement of pores
in tangential section, but Taylor's description (1910 : 132) and Bedford's figure (1936,
fig. 69) confirm my observations from transverse sections which suggest the pores are
small, isolated and without any important modifications in the lateral bulges. No
vesicular tissue.
DIMENSIONS
Cup: (mm.)
Diameter ....... about 85
Intervallum
without bulges ...... 8
largest part . . . . . . 19
Interseptum . . . . . . 1-07
Synapticulae
horizontal ...... 1-2
vertical ....... 0-6-2
Outer wall:
No. of pores per interseptum. ... 4
Diameter of pores . . . . . 0-13
Partitions . . . . . . • 0-06
Inner wall :
No. of pores per interseptum .... i
Diameter of pores . . . . . ° ' 33
Horizontal partitions . . . . . 0-67
Thickness 0-67
Taeniae :
Thickness . . . . . . . o-i
DISCUSSION. This is the largest species of Flindersicyathus (Pycnoidocyathus)
known. It has regular annular bulges, the skeletal tissue of the wavy taeniae
occupies a greater area than the pores, and the synapticulae are more numerous than
in any other species of this genus.
352 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
Flindersicyathus (Pycnoidocyathus) simplex (Taylor)
(PL 15, fig. 2)
1910 Pycnoidocyathus simplex Taylor : 134, pi. 2, fig. 70.
1936 Pycnoidocyathus simplex Taylor; R. & W. R. Bedford : 15, pi. 15, fig. 70.
HOLOTYPE. Probably in the South Australian Museum.
MATERIAL. B.M. (N.H.) S 4824.
DESCRIPTION. Broken piece showing well-preserved inner wall. Taeniae are
radial, not waved and are joined with the synapticulae, chiefly towards the exterior.
Outer wall is only see on part of a transverse section, where a small weathered-out
surface shows that the pores are arranged in irregular quincunx. Inner wall has a
vertical row of pores per intersept, each pore having a raised lower edge and is
vertically elongated but flattens towards the top. The rows of pores alternate in
quincunx.
DIMENSIONS
(mm.)
Cup:
Height (pars) ..... 40
Diameter ...... about 22
Intervallum . . . . . 5-5
Central cavity ..... 13
Intervallum coefficient . . . 0-4
Intertaenial space .... i
Outer wall :
Diameter of pores .... 0-40-0-33 (hor.)
Skeletal partitions .... 0-20-0-20
Thickness . . . . . 0-13
Inner wall:
Diameter of pores .... i -69-1 -42 (hor.)
Vertical partitions . . 0-20
Horizontal partitions . o • 47
Taeniae :
Diameter of pores ... o«6
Vertical partitions . . . . 0-13
Horizontal partitions . . . 0-20
Thickness . . . . . o • i
DISCUSSION. The measurements of the specimen described above correspond to
those given by Taylor (1910). R. & W. R. Bedford (1936 : 15, fig. 70) pointed out
and sketched the regular annular bulges from an example of the same species. F. (P.)
simplex differs from typical Pycnoidocyathus by the scarcity of its synapticulae and
the straight taeniae.
Flindersicyathus (Pycnoidocyathus) maximipora (R. & W. R. Bedford)
(PI. 15, fig. 3)
1934 Pycnoidocyathus maximipora R. & W. R. Bedford : 3, pi. 2, figs. ga-c.
1936 Pycnoidocyathus maximipora R. & W. R. Bedford : 15, pi. 15, fig. 71.
HOLOTYPE. B.M. (N.H.) S 4150.
LOWER CAMBRIAN ARCH AEOC Y ATM A FROM AUSTRALIA 353
DESCRIPTION. One broken specimen with a single annulate bulge. Intervallum
crossed by un-waved radial, porous taeniae, with the skeletal tissue forming a greater
area than the pores, and sparse synapticulae, that occur mainly in the lower part of
the fragment. The taenial pores, are round and arranged in lines, which curve
upwards and outwards, near the inner wall; but are oval and bigger near the outer
wall and inside the annular bulge. The thin, irregular, outer wall is strengthened by
bars springing from the taeniae, but both partial weathering and preservation give
an irregular and false idea of its original structure.
The inner wall is composed of very short piled-up pipes, leading obliquely upward,
with one row per intertaenial space, which alternates with the next. The cross-sec-
tion of the pipe is vertically oval. Each pipe is withdrawn from the one immediately
above and its sides are stretched slightly to join with its neighbours. Thus, the
inner wall does not present a flat surface to the central cavity but has the appearance
of a rasp. The various sections have suggested to previous authors that there is a
louvre-like plate at the lower part of each pore, but, in fact, these are the short pipes.
DIMENSIONS
(mm.)
Cup:
Height (pars) 35
Diameter 22
Intervallum 6
Intertaenial space i • 3
Outer wall:
Diameter of the pores o • 4
Partitions o • 3
Thickness o • 4
Inner wall:
Diameter of the pores i -49-2 ' 71
Lateral partitions 0-13
Horizontal partitions 2 • 03
Taeniae :
Diameter of the pores near the inner wall o • 6
near the outer wall o • 8-2 • 35
Partitions 0-27-0-40
DISCUSSION. This species differs from others by the greater development of
tubes in the inner wall and its relatively narrower taenial pores.
F Under sicyathus (Pycnoidocyathus) vicinisepta (R. & W. R. Bedford)
(PI. 15, fig. 4)
1936 Pycnoidocyathus vicinisepta R. & W. R. Bedford : 15, pi. 16, fig. 72.
HOLOTYPE. Probably in the South Australian Museum.
MATERIAL. B.M. (N.H.) S 4825.
DESCRIPTION. A longitudinally cut fragment. Several annulate horizontal bulges
are visible, they are rather narrow and close together. Radial un-waved taeniae
354 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
joined by numerous synapticulae. In the bulges, the taeniae are waved with opposite
crests connected by synapticulae, so that the intervallum has the aspect of a polygonal
mesh. A very small part of the outer wall is preserved in which the pores are irregular
and polygonal. The inner wall has one vertical row of pore-tubes leading up into the
central cavity, in each intertaenial space. The rows alternate.
DIMENSIONS
(mm.)
Cup:
Height (pars — in two pieces) 95
Diameter 22
Bulge 4
Intervallum without bulge 4
Central cavity 14
Intertaenial space o • 33
Intersynapticular space o • 47-0 • 67
Outer wall :
Diameter of pores o • 06
Skeletal tissue o • 05
Thickness not known
Inner wall :
No. of rows of pores per intertaenial space i
Horizontal partitions 0-27
Vertical partitions o • 40
Diameter of pores o • 40
Thickness 0-40
Taeniae :
Diameter of pores 0-27
Horizontal partitions 0-81
Vertical partitions o • 67
Thickness o-io
DISCUSSION. As far as I can tell, after measuring the drawings of Bedford, the
coefficients correspond to those of vicinisepta. This species differs from other
Pycnoidocyathus in having many more taeniae.
Family METAGYATHIDAE R. & W. R. Bedford, 1934
DIAGNOSIS. Cups growing from an apex without central cavity, occupied by rods,
plates and dissepiments. Adult stages with more or less definite radial septa, the
pores of which are arranged in rows inclined upwards and outwards from inner to
outer wall. Dissepiments and sometimes synapticulae present. Simple porous
outer and inner walls, with the pores sometimes screened by a microporous sheath or
pellis.
COMPOSITION OF THE FAMILY. Protopharetra Bornemann 1887, Volvacyathus
Debrenne 1961, Dendrocyathus Okulitch & Roots 1947, ? Shidertycyathus Krasnopeeva
1959, Metaldetes Taylor 1910, Cambrocyathus Okulitch 1937, Cambrocyathellus
Zhuravleva 1960, Okulitchicyathus Zhuravleva 1960, Pamnacyathus R. & J. Bedford
1937, ? Ardrossacyathus R. & J. Bedford 1937, Metafungia R. & W. R. Bedford 1934.
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA 355
Genus METALDETES Taylor, 1910
1910 Metaldetes Taylor : 151, pi. 15, figs. 86-88, t.-figs. n, 37 and 38.
1934 Metacyathus R. & W. R. Bedford : 5.
?i957 Bedfordcyathus Vologdin : 182 and 209.
TYPE SPECIES. By monotypy Metaldetes cylindricus Taylor 1910, the holotype of
which is in the University of Adelaide.
DIAGNOSIS. Solitary, or sometimes colonial cups; the central cavity and the inner
wall slowly develop later. In the lower part of the cup, the irregularly arranged
skeletal structures (bars, rods, plates and vesicular tissue) fill the whole inner space.
Much later, the central cavity is defined by the formation of a regularly porous inner
wall and the structures of the intervallum are arranged in radial plates ; the tangential
links (synapticulae and dissepiments) may continue into the adult stage.
Unfortunately, the type specimen of the genus has not been re-described yet and
one has to rely on the original account and illustrations. According to Taylor
(1910 : 151), the rugose character of the outer wall causes a transverse section to
resemble a series of tridents. R. & W. R. Bedford (1934 : 5) considered the outer wall
to be a double porous sheath, and my own observations agree with this (Debrenne
1964 : 219). Hill (1965 : 119) mentions that the longitudinal ribs between the rows
of pores are connected together, some distance behind the outer edge, by transverse
bars. The inner wall, which is absent in the lower part of the cup, was described by
Taylor (1910 : 152) as being " strongly ridged on its septal side ". These ridges
correspond to the beginnings of the taeniae, which are slightly thickened towards the
inner wall. The number of pores per intersept is not certain (i, 2, or 3), nor their
shape and size.
DISCUSSION. A complete revision of the type material is desirable, but although
Metaldetes is incompletely known, it is possible to regard two subsequent genera,
Metacyathus Bedford and Bedfordcyathus Vologdin, both found at the same locality
and level, as synonyms. Okulitch (1955 : E 16), Zhuravleva (1960 : 283) and myself
(Debrenne 1964 : 220) have previously considered Metacyathus a synonym of Metal-
detes, but Vologdin (1957 : 43) and Hill (1965 : 118) thought it to be a separate genus.
R. & W. R. Bedford established the genus to include those species in which the taeniae
occur as straight radial septa in the inner two-thirds of the intervallum, but are
separate from the outer wall, as in the type M . taylori R. & W. R. Bedford
Examination of the holotype of M . taylori shows that the structures of the inter-
vallum are disturbed by the occurrence of exothecal lamellae, but that, at other
levels, the plates continue from one wall to the other. The porosity of the outer wall
depends on the presence or absence of the exotheca; the inner wall has one vertical
row of pores per intertaenial space in the lower part of the cup, but two or more in
the upper part, with probably a second wall.
Bedfordcyathus was established by Vologdin, after he had studied Bedford's figures,
to group into a separate genus those species that have a strong development of vesi-
cular tissue. The holotype of the type species (M. irregularis Bedford), now in the
British Museum (Natural History), is described below. It shows that there is no
356 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
appreciable difference from Metacyathus, in the structure of the intervallum (apart
from the abundance of vesicular tissue), or in that of the walls. I think that vesicular
tissue is too inconstant and variable a character to be sufficient grounds for establish-
ing a new genus.
The genus Metaldetes is therefore denned as consisting of cups which grow from a
base that does not have a distinct central cavity, where the skeletal elements have
not attained a clear radial arrangement, and where tangential links (synapticulae and
vesicular tissue) are numerous. Subsequently, the central cavity is differentiated by
a more definite development of the inner wall, which is, at first, part of the intervallum
(with one pore per intertaenial space), but then becomes a more distinct structure
(with two pores per intertaenial space) and is, perhaps, protected on its central cavity
side by a microporous sheath or vesicular membrane. At this same level, the inter-
vallar elements generally form strong, compact, radial taeniae with high oval pores
the diameter of which increases along upwardly curving lines from the inner to the
outer wall; the largest pores being close to the outer wall. It is in this region that
positional changes and external influences occur, the septa become less regular and
sometimes revert to curved taeniea. The synapticular links may persist but generally
disappear in the adult stage, whereas vesicular tissue is still abundant. The outer
wall has a basic layer with irregular polygonal pores, overlapped by a second micro-
porous sheath, which disappears when the cup is surrounded by exothecal lamellae.
Fossils reach considerable size.
RELATIONSHIPS AND DIFFERENCES
Volvacyathus Debrenne 1961 has two distinct stages, the apex being quite different
from the adult. The walls are still connected with the intervallar network, the septa
are not clearly denned and their pores are wider than the intervening skeletal tissue.
This genus is also close to Protopharetra Bornemann 1887.
Cambrocyathus Okulitch 1937 closely resembles Metaldetes in having abundant
vesicular tissue, but the adult stage is reached more quickly. The central cavity is
not free and is filled by thin skeletal rods and vesicular tissue, which persists as a
pellis to the top of the cup. These structures are also found as an exotheca. The
well-developed taeniae generally have a laminated structure and are perforated by
pores, that are almost in horizontal lines. Scarce synapticulae or branching septa,
but abundant vesicular tissue, which earlier authors have interpreted as true synap-
ticulae. Reference material on loan from the Geological Survey of Canada and a
topotype from the Yale Peabody Museum showed these features (Text-fig 14).
Septa of Cambrocyathus-type (see Debrenne 1964 : 88) are the final stage in the
process of radial partition. This starts with Protopharetra, leads through Volvacya-
thus (see Debrenne 1964, fig. 51) and the first Metaldetes (proteiis and dissutus), with
their broken septa, to the complete septa of Metaldetes irregularis and finally, to the
regular septa of Cambrocyathus.
COMPOSITION OF THE GENUS. M. cylindricus Taylor 1910, M. columbianus
(Okulitch 1943), M. dissepimentalis (Taylor 1910), M. dissutus Debrenne 1964, M.
irregularis (R. & W. R. Bedford 1934), M. ramulosus R. & J. Bedford 1937, M. solidus
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA 357
FIG. 14. Cambrocyathus profundus (Billings) X4
358
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
(Okulitch 1957), M. spiralis R. & W. R. Bedford 1936, M. superbus R. & W. R.
Bedford 1936, M. taylori (R. & W. R. Bedford 1934), M. proteus (Bornemann 1887).
Metaldetes dissepimentalis (Taylor)
(PI. 16, figs. 1-3)
1910 Archaeocyathus dissepimentalis Taylor : 128, pi. 10, fig. 53.
1934 Metaldetes conicus R. & W. R. Bedford : 5, figs 26, 28 and 31.
1936 Metaldetes conicus R. & W. R. Bedford : 18, pi. 18, fig. 77.
HOLOTYPE. Possibly at Adelaide University.
MATERIAL. B.M. (N.H.) S 4176-4182.
DESCRIPTION. Large conical cups. In the first stages, the central cavity is not
formed and the entire cup is occupied by un-orientated taeniae, which are joined by
numerous dissepiments that are arranged as horizontal vesicles. Much later, the
cavity develops, the dissepiments become less abundant and the taeniae form regular
radial plates. The general aspect recalls that of Cambrocyathus profundus Billings.
The outer wall is twofold: a basal wall covered externally by a very thin micro-
porous sheath. The inner wall is also double; the main wall having two rows of
large irregular pores per intersept, with the diameter of the pores being much greater
than the width of the skeletal tissue between them; with a microporous sheath,
similar to that of the outer wall, occurring on the central cavity side. The micro-
porous sheaths are not well known for they are often eroded and no good tangential
sections have been seen. Consequently, the arrangement, size and shape of the
micropores is not certain, they appear to be similar to those of the secondary wall in
Metafungia.
DIMENSIONS
Cup:
Height (pars)
Diameter
Intervallum
Interseptum
Intervallum coefficient
Outer wall:
No. of rows of pores per
interseptum
Diameter
Skeletal partitions
Micropores: diameter
skeletal partitions
Thickness
S 4176
(mm.)
60
24
6
0-4-0-7
S 4178
(mm.)
13
4
o-5
0-8
S 4179 S 4180
S 4180 up.
low. S 4181
(mm.)
(mm.
13
0-7
15
21
6
0-6
S 4182
(mm.)
40
45
7'5
i-35
0-25
dissepimentalis
Taylor
(mm.)
22
6
7
0-6
I-
-3
1-3
2
2
1-3
2
0
• 17
0'33
O
•2
O
•20
O'
•25-0
•40
O
•30
0
•30
O-2O
O
•15
O
•15
O'
•15-0
•20
O
•20
0
°3
O
•06
O
•O6
.
.
o
•06
O
06
. .
.
.
O-2O
0-20
LOWER CAMBRIAN ARCHAEOCYATH A FROM AUSTRALIA
DIMENSIONS — continued
359
Inner wall:
No. of pore rows per
interseptum
Diameter
Skeletal partitions
Micropores: diameter
skeletal partitions
Thickness
Septa :
Diameter
Skeletal partitions
Thickness
S 4176
S 4178
84179
S 4180
low.
S 4180
up.
S 4181
(mm.)
(mm.)
(mm.)
(mm.)
2
2
••
i-3
0-67
0-16
0-47
0-16
••
o-io
0-40
0-13
0-03
0-03
0-13
S 4182
(mm.)
1-2
o-33
o-io
0-03
0-03
0-40
o • 40-0 • 60
O-2O
dissepimentalis
Taylor
(mm.)
1-2
0-30
o-io
0-03
0-03
DISCUSSION. The description and figures given by Taylor for Archaeocyathm
dissepimentalis correspond with measurements taken from the photographs of
Metaldetes conicus Bedford. The two species are, therefore, considered to be synony-
mous and dissepimentalis, the older, is the name conserved.
Metaldetes irregularis (R. & W. R. Bedford)
(PL 16, fig. 4)
1934 Metacyathus irregularis R. & W. R. Bedford : 6, pi. 5, fig. 29.
1957 Bedfordcyathus irregularis (Bedford) Vologdin : 43.
1964 Metacyathus-Bedfordcyathus irregularis (R. & W. R. Bedford); Debrenne : 220 and 231.
1965 Bedfordcyathus irregularis (R. & W. R. Bedford); Hill : 118, fig. 22.9.
It has not been possible to locate the original description by Vologdin of his genus
Bedfordcyathus. Both Debrenne (1964) and Vologdin (1957, 1962) give the year 1955
for this genus, but, unfortunately, there is no mention of Bedfordcyathus in the paper
cited under that year, in either bibliography (Dokl. Akad. Nauk SSSR, 103, i). In
subsequent papers, Vologdin gives the year as 1956, but the only paper available for
that year merely includes the genus in a list showing the classification of the Archaeo-
cyatha. This paper appeared in a slightly more elaborate form during 1957, in two
journals: Ada paleont sinica, 5: 173-222 and Annls. Cent. Etud. Docum paleont.,
23 : 33-80 and it is the latter, which is taken as the basis of the date given in this
paper and that of Hill (1965). However, it would appear that the paper referred to,
is included in the yet unpublished Vol. 3 of " El sistema Cambrico . . . ", Int. Geol.
Congr. Mexico, 1955, quoted in some references as 1961.
HOLOTYPE. B.M. (N.H.) S 4189, by monotypy.
OTHER MATERIAL. Paratypes B.M. (N.H.) S 4188, S 4190.
DESCRIPTION. Conical cup, with large irregularly- waved intervallum. Double
outer wall; first, a thick basal one, pierced by funnel-shaped pores that widen towards
GEOL. 17, 7
28
360 LOWER CAMBRIAN ARCH AEOCYATH A FROM AUSTRALIA
the exterior ; this is covered externally by a very thin microporous sheath. The inner
wall is not well known, but from a small poorly preserved surface, appears to be
covered by an irregular, microporous, thin wall, which screens the simple pores of the
intervallum.
The taeniae in the upper part, are straight, radial and completely cross the inter-
vallum. In longitudinal section, the pores appear circular and alternate in regular
lines, curving shallowly towards the exterior from the inner to the outer wall. The
skeletal tissue and the pores, occupy equal proportions of the total surface area.
Considerable development of vesicular tissue, formed of imbricate, horizontally
elongated vesicles, that cross several loculi, but do not penetrate into the central
cavity.
DIMENSIONS
(mm.)
Cup:
Height (pars) 92
Diameter about 35
Intervallum 10
Intervallum coefficient 0-6
Interseptum i'35
Outer wall :
No. of rows of pores per interseptum 2
Diameter of pores 0-27
Partitions 0-20-0-40
Micropores o-io
Inner wall:
Diameter of pores o • 35-0 • 67
Partitions 0-13
Micropores 4 to each main pore
Diameter of micropores o • 05
Thickness o • 06
Taeniae :
Diameter of pores o • 33-0 • 54
Partitions 0-27-0-33
DISCUSSION. The strong development of vesicular tissue and the absence of exo-
thecal lamellae, are the main differences between this species and others in the genus.
Metaldetes taylori (R. & W. R. Bedford)
(PI. 13, figs. 1-3)
1934 Metacyathus taylori R. & W. R. Bedford : 5, pi. 5, fig. 10.
1936 Metacyathus taylori R. & W. R. Bedford; R. & W. R. Bedford : 18, pi. 18, fig. 80.
1964 Metaldetes taylori (R. & W. R. Bedford) Debrenne : 220.
1965 Metacyathus taylori R. & W. R. Bedford; Hill : 118, pi. 9, figs. 2a-d.
HOLOTYPE. By monotypy, B.M. (N.H.) S 4185-4187 (three pieces of the same
specimen) .
DESCRIPTION. Although the three pieces are said to come from the same cup, it is
no longer possible to prove this with any certainty, for having been subjected to
various palaeontological techniques (cutting, etching etc.), they do not fit together.
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
361
Nevertheless, there is little chance of the lower part belonging to another specimen.
This is pointed out because the lower part has some structures obviously different
from the other two pieces and these are important for generic determination.
The lower part is conical ; radial, slightly porous taeniae are connected by synapti-
culae, which develop between the pores. The inner wall has a single row of pores per
interseptum. The outer wall is well denned and separates the intervallum from the
exothecal lamellae. Dissepiments present.
In the middle and upper parts of the cup, the inner wall has two rows of pores per
interseptum; the outer wall is not yet separate from the exothecal structures, which
disturb the outer part of the intervallum and give the septa the form of curved taeniae.
No synapticulae, but dissepiments persist. The pseudo-septa are mainly radial at
the inner and middle parts of the intervallum, but are interrupted towards the outer
wall and disappear near the exothecal lamellae.
DIMENSIONS
84185
Apex
Cup:
Height (pars)
Diameter
Intervallum
Intervallum coefficient
Interseptum
Outer wall :
Thickness
Inner wall :
No. of pore rows per interseptum
Diameter
Skeletal partitions
Thickness
Taeniae
Diameter of pores
Thickness
(mm.)
35
16
0-72
0-06
S 4186
(mm.)
55
35
24
ii
7
O'
o
upper
lower
upper
lower
upper
lower
S 4187
(mm.)
40
35
disappeared
2
o-33
O-2O
?
o • 40-0 • 60
0-40
0-27
disappeared
2
o-33
O-2O
?
O-6o
0-40
O-25
DISCUSSION. Metaldetes taylori (R. & W. R. Bedford) differs from M. irregular-is
(R. & W. R. Bedford) by its narrow central cavity, weak development of vesicular
tissue and the abundance of exothecal lamellae. It is little different from M.
dissepimentalis (Taylor), which has straighter septa, a well-defined, double-porous
outer wall, but is without any exothecal lamellae.
Genus METAFUNGIA R. & W. R. Bedford, 1934
1934 Metafungia R. & W. R. Bedford : 5.
1964 Metafungia R. & W. R. Bedford; Debrenne : 219
1965 Metafungia R. & W. R. Bedford; Hill : 119.
TYPE SPECIES. Metafungia reticulata R. & W. R. Bedford, by monotypy.
GEOL. 17, 7 '*
362 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
DIAGNOSIS. Cup with central cavity that becomes free of skeletal tissue late in its
development. The walls are double-porous. The intervallum is crossed by straight
taeniae, which have their pores arranged in lines curving upwards and outwards;
numerous synapticulae join the taeniae. The vesicular tissue is present at the base
and in contact with tersioid outgrowths.
COMPOSITION OF THE GENUS. Metafungia reticulata R. & W. R. Bedford.
Metafungia reticulata R. & W. R. Bedford
(PL 18, figs. 1-3)
1934 Metafungia reticulata R. & W. R. Bedford : 5, figs. 23 a-e.
1965 Metafungia reticulata R. & W. R. Bedford; Hill : 119, pi. 10, fig. 2 and text-fig. 22, 12 a-d.
HOLOTYPE. B.M. (N.H.) S 4184. Other Material S 4183.
DESCRIPTION. Large conical cup, the apex is surrounded by exothecal tersioid
outgrowths and the central cavity is filled with endothecal tissue. The intervallum is
crossed by straight, radial taeniae, which are perforated by pores, aligned upwards
and outwards, in oblique rows. The diameter of these pores increases from the inner
to the outer wall, giving an irregular appearance. The numerous synapticulae con-
nect at the skeletal junctions of the taeniae. The vesicular tissue is only developed in
the exo- and endothecal regions. The intervallar mesh, with some development of
skeletal tissue and vertical closing of the apertures, corresponds to an outer wall. A
regular micro-porous thin wall covers this externally. The inner wall may also be
interpreted as a double wall. The inner part of each taenia has pores no larger than
0*7 mm; these taeniae are joined by small plates, resembling widened synapticulae, to
form a small tube. On the central cavity side, a thin sheath is developed, with two
pores to each tube, and has a more or less regular, hexagonal appearance.
DIMENSIONS
S 4183 Base S 4184 Top
(mm.) (mm.) (mm.)
Cup:
Height .. 35
Diameter 12 . . 21
Intervallum 2-5 . . 5
Intervallum coefficient 0-3 . . 0-4
Interseptum 0-8 .. i-oi
Synapticulae : radial distance . . . . o • 61
vertical distance . . . . o • 61
Outer wall:
Diameter of main pores . . 0-70
Partitions .. 0-27
Thickness . . 0-2
Micropores : diameter . . 0-07
partitions . . 0-06
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
363
DIMENSIONS — continued
Inner wall :
Diameter of tubes
Partitions
Length of tubes
Inner pores : diameter
partitions
Septa :
Thickness
Diameter of inner pores
Diameter of outer pores
Synapticulae :
Thickness
S 4183 Base
(mm.)
(mm.)
o • 7-0 • 7
0-27
0-70
O-2O
0-13
0-I3
O-25
0-40
0-13
S 4184 Top
(mm.)
DISCUSSION. Metafungia reticulata has an intervallar structure similar to that of
Flindersicyathus graphicus (R. & W. R. Bedford). However, its size is much greater
and the connecting synapticulae do not occur at every skeletal junction, but the
essential difference is in the structure of the walls.
Family METAGOSCINIDAE R. & W. R. Bedford
DIAGNOSIS. Two-walled porous cups. Taeniae and sparse convex tabulae.
COMPOSITION OF THE FAMILY. Metacoscinus R. & W. R. Bedford 1934, Pycnoido-
coscinus R. & W. R. Bedford 1936, Paracoscinus R. & W. R. Bedford 1936, Claruscya-
thus Vologdin 1932, Gabrielsocyathus Debrenne 1964.
Genus METACOSCINUS R. & W. R. Bedford
1934 Metacoscinus R. & W. R. Bedford : 6, pi. 5, fig. 27.
1965 Metacoscinus R. & W. R. Bedford; Hill : 133, pi. 12, fig. i, text-fig. 5 a-d.
TYPE SPECIES. Metacoscinus reteseptatus R. & W. R. Bedford, by monotypy.
DESCRIPTION. Conical cups, slightly waved externally. Oblique and vertical
rods form the skeletal tissue of taeniae crossing the intervallum. These rods have
considerable openings between them, which are much taller than wide and are
roughly polygonal in shape. The outer pores are the largest. Less frequent horizon-
tal structures regarded as tabulae, are perforated by small circular pores, that are
separated by " linteaux " of varying size, and bear little irregular tubercles on each
side.
Outer wall has funnel-shaped pores, circular on the intervallum side, larger and
more irregular on their open outer side. Thin inner wall with two rows of large
polygonal pores per intersept, separated by a thin skeletal mesh. R. & W. R.
Bedford figured a specimen (1936 : fig. 83), which had several tubercles on its central
cavity side.
364 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
COMPOSITION OF THE GENUS. Metacoscinus reteseptatus R. & W. R. Bedford 1934,
? Metacoscinus insigne R. & W. R. Bedford 1936.
DISCUSSION. Metacoscinus insigne is retained in this genus with some misgiving,
until the type material can be revised. It has no visible tabulae and a thick outer
wall with labyrinthoid pores. Nothing is known about the structure of these pores,
whether their distorted shape is due to the fusion of several tubercles, as seen in the
tabulae of M . reteseptatus, or results from lateral connection between neighbouring
pore-tubes.
I have had the opportunity to examine the type material of Metacoscinus gabriel-
sensis (Okulitch 1955), M. deasensis (Okulitch 1955) and M. poolensis (Kawase &
Okulitch 1957) loaned by the Geological Survey of Canada and have already suggested
(Debrenne 1964 : 248), that these species should be regarded as a separate genus,
Gabrielsocyathus Debrenne 1964, with G. gabrielsensis (Okulitch) as type species. This
differs from Metacoscinus in having simply-porous, regularly-spaced tabulae ; branch-
ing, or waved porous taeniae ; a double, porous, outer wall with exotheca ; endothecal
lamellae partially filling the central cavity and extremely abundant vesicular tissue.
The species poolensis (Kawase & Okulitch) is included in Gabrielsocyathus with some
reservations. The septa appear to undulate and bifurcate and have fewer pores than
the other two species; the plates thicken at the junction with the septa; no exotheca
occurs and the vesicular tissue is less abundant. The poor preservation of the speci-
men prevents confirmation of the presence of fine, irregular pores in all the skeletal
plates.
Zhuravleva (1960 : 311) remarks that Paracoscinus and Metacoscinus are very
close to each other and goes on to suggest that they might represent one genus, but
since she had no material, it was difficult to make a definite decision. I have not seen
the type of Paracoscinus, but from the figures and descriptions of R. & W. R. Bedford
(1936 : 18, pi. 20, figs. 85-86), I consider that it differs from Metacoscinus in its wall
structure (microporous and not a large open mesh) ; the abundant and regular plates,
which have polygonal pores and are without tubercles ; the thin outer wall and the
inner wall with a single row of pores per interseptum.
Metacoscinus reteseptatus R. & W. R. Bedford
(PL 17, figs. 1-4)
1910 Archaeocyathus retesepta Taylor : 120, pi. 7, figs. 39-40.
1934 Metacoscinus reteseptatus R. & W. R. Bedford : 6, pi. 5, fig. 27.
1936 Metacoscinus reteseptatum R. & W. R. Bedford; R. & W. R. Bedford; 18, pi. 19, fig. 83.
1965 Metacoscinus reteseptatus R. & W. R. Bedford; Hill : 132-3, pi. 12, fig. i, text-fig. 24, 5 a-d.
MATERIAL. Syntypes B.M. (N.H.) S 4191-4195. Other material S 4762, S 4772,
S 4743, S 7633.
DESCRIPTION. Conical cups with transverse annular bulges. Outer wall has two
vertical rows of funnel-shaped pores per intersept, the larger aperture on the outside,
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
365
with an irregular rim. Inner wall, also, has two vertical rows of pores, but these are
polygonal and separated by very thin skeletal rods, without tubercles. Regular,
radial pseudosepta, perforated by numerous hexagonal pores that are elongated
upwards, or upwards and outwards in the annular bulges. Little vesicular tissue
except in the lower part, where the pseudosepta are not so well developed and skeletal
elements fill both the central cavity and the intervallum. Sparse tabulae, which have
tubercles occurring between their two rows of quincunxial pores per intersept. The
regular pattern of the pores is disturbed by the coalescence of neighbouring pores, or
by anastomosed tubercles, giving these horizontal plates a vermiculate aspect, when
seen from above.
DIMENSIONS
Cup:
Height (pars)
Diameter
Intervallum
Central cavity
Interseptum
Intertabulum
Outer wall :
No. of pore rows
per interseptum
Diameter
Vertical partitions
Horizontal partitions
Thickness
Inner wall :
No. of pore rows
interseptum
Diameter
Vertical partitions
Horizontal partitions
Thickness
Septa :
No. of pore rows
Diameter
Vertical partitions
Horizontal partitions
Thickness
Tabulae
No. of pore rows
Diameter
Vertical partitions
Horizontal partitions
Thickness
S 4191
(mm.)
21
12
2
8
i-y-3'4
2-3
0-15
0-06-0-10
0-06-0- 10
0-30
0-07
0-07
O-2O
0-15^-37
0-07
O-II
2
O-II
o- ii
O-II
0-15
S 4192 &
S 4193
low
(mm.)
base
7
full
O-22
0-26
S 4193
upper
(mm.)
10
2
4
0-26
2
0-07
O-O7
O-II
6
O • 22-O • 40
0-07
0-07
S 4195
(mm.)
25
25-30
6
ii
o-75
2
0-18
0-15
0-15
0-18
2-3
0-18-0-37
0-07
0-07
O-II
0-56-1-51
O-26-O-52
0-18
0-18
o-ii
84762
(mm.)
20
15
2-45
unknown
0-22
0-15
0-06-0-10
o-io
unknown
1-2
0-22-0-30
0-07
0-15
unknown
unknown
0-26
0-13
0-07
366
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
DIMENSIONS — continued
Cup:
Height (pars)
Diameter
Intervallum
Central cavity
Interseptum
Intertabulum
Outer wall:
No. of pore rows
per interseptum
Diameter
Vertical partitions
Horizontal partitions
Thickness
Inner wall:
No. of pore rows
interseptum
Diameter
Vertical partitions
Horizontal partitions
Thickness
Septa :
No. of pore rows
Diameter
Vertical partitions
Horizontal partitions
Thickness
Tabulae :
No. of pore rows
Diameter
Vertical partitions
Horizontal partitions
Thickness
84772
(mm.)
unknown
16
3'5
S4743
(mm.)
12
10
2
O-22
2-3
0-13
0-07
O- II
unseen
$4743
lower
(mm.)
84194
lower
(mm.)
84194
upper
(mm.)
7
30
9
16
i'35
4 -3 clear
i-45
skel. el.
4-8
o-33
0-41
o-37
unseen 2 2
0-26-0-15 0-26-0-26
0-05 0-26—0- i 8
o-n
0-18
2 unseen unseen
unseen . . 0-18
0-07
0-07 . . o-ii
o-n
o-io
not visible
0-15
87633
(mm.)
15
4
o-n
2-3
0-15
O-II
O-II
7
7
5
7
o-45
o- 13-0-22
0-13
0-49
0-34-1-7
0-26
O-II
0-07
0-07
0-07
0-15
O-II
0-15
0-15
0-15
O-II
0-07
O-II
O-II
O-II
retesepta
Taylor
(mm.)
90
25
2
O-2O
O-II
2-3
0-15
o-io
0-10
0-15
0-15
DISCUSSION. R. & W. R. Bedford (1934 : 6) compared this species with Archaeo-
cyathus retesepta Taylor, noting the likeness between the taeniae, but no horizontal
structures were known in A. retesepta. Nevertheless, from Taylor's figures (PL 7,
figs. 39 and 40), one can perhaps consider that the structures normal to those in the
general direction are the beginnings of tabulae. Amongst all the specimens in the
Bedford collection at the British Museum (Nat. Hist.), only S 4195 and S 4194 show
these rare horizontal structures. The dimensions of the two species are of the same
order and they are probably synonymous. Before joining them as a single species we
need more material, in order to know the frequency and significance of these horizon-
tal structures and to decide whether they have any systematic value, or are true
tabulae, or are accidental features.
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 367
? Family METAGOSCINIDAE R. & W. R. Bedford, 1936
Genus PYCNOIDOCOSCINUS R. & W. R. Bedford, 1936
1936 Pycnoidocoscinus R. & W. R. Bedford : 19, pi. 20, fig. 87.
TYPE SPECIES. Pycnoidocoscinus pycnoideum R. & W. R. Bedford, by original
designation.
DIAGNOSIS. Cups with annular bulges on the outer wall, with the inner wall
remaining cylindrical. Radial septa and arched tabulae. The outer wall is a porous
mesh; the inner wall with both horizontal and vertical rows of rectangular pores.
DISCUSSION. As the early stages are not known, it is difficult to classify this genus
in either the Irregularia — indicated by the irregular outer wall and septa with
numerous pores, or amongst the Regularia — as suggested from the type of inner wall
with its rectangular apertures, the tabulae with the axis of curvature inside the
intervallum, or the pore pattern of Retecoscinus-type. Only having examined a
single broken fragment, it is not possible for me to offer any solution and the genus
Pycnoidocoscinus is retained in the Metacoscinidae, but with some reservations.
COMPOSITION OF THE GENUS. Pycnoidocoscinus pycnoideum R. & W. R. Bedford.
Pycnoidocoscinus pycnoideum R. & W. R. Bedford
(PI. 18, figs. 4-8)
1936 Pycnoidocoscinus pycnoideum R. & W. R. Bedford : 19, pi. 20, fig. 87.
MATERIAL. B.M. (N.H.) S 4832. A syntype P 990 is in the South Australian
Museum, Adelaide.
DESCRIPTION. Only one weathered and poorly preserved specimen has been
examined. The intervallum is crossed by radial septa and arched tabulae. The
septa are thickened near the inner wall and sometimes near the outer wall, but are
very thin in the middle. Septal pores are not visible. Pores on a small, tangential
surface of a tabula resemble those of Retecoscinus Zhuravleva (1960), with two
alternating rows of oval to rectangular pores.
Outer wall eroded away; R. & W. R. Bedford have described it as " a layer of fine
pores supported by an irregular mesh ". Inner wall has two rows of pores per inter-
septum and two rows of pores per intertabulum.
The vertical partition between the two interseptal pores, is the beginning of a new
septum, which only develops in the inner third of the intervallum. Sometimes
opposite this crude septa, another arises from the outer wall to meet it.
DIMENSIONS
(mm.)
Cup:
Height (pars) 30
Diameter 22
Interseptum (at the inner wall side) o • 33
Intertabulum 1-22
368 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
DIMENSIONS — continued
(mm.)
Outer wall: weathered away
Inner wall :
Diameter of pores 0-33
Vertical partitions o • i
Horizontal partitions o • 06
Septa
Diameter of pores o • 06
Vertical partitions o • 2
Thickness: middle 0-06
near inner wall o • 2
Tabulae :
No. of pores per loculus 2 x 22
Diameter of pores o • 27-0 • i
Partitions 0-1-0-13
Thickness o • i
DISCUSSION. The rows of rectangular pores of the inner wall and the presence of
" rods " in the middle of the interseptum (interpreted here as the beginnings of new
septa) are good reasons for including S 4832 in P. pycnoideum. Unfortunately, poor
preservation and the lack of an apex prevents any clarification of the systematic
position of Pycnoidocoscinus.
III. STRATIGRAPHICAL CONCLUSIONS
The excellent silicified fossils found in the Australian Cambrian, as well as being
extremely useful for anatomical studies of the Archaeocyatha, are also of considerable
stratigraphical interest.
We have little information on the precise horizon in the Ajax Limestone, from which
the Bedfords collected their fossils. According to Daily (1956 : 129), ". . . Most, if
not all, of the species described from the Ajax Mine area by Taylor and the Bedfords,
are believed to have come from beds containing this assemblage [i.e. faunal assem-
blage No. i], for it is significant that no fossils characteristic of faunal assemblage No.
2 have been reported by them. Such fossils would almost certainly have been noticed
if they were present . . . ". However, Walter (1967) comments (p. 145) that the
rich Ajax fauna described by the Bedfords & Taylor, cannot be placed in Daily's
scheme of faunal units. This is because, in an area of complex structure, it was not
collected in sequence and the additional evidence of the fauna associated with the
archaeocyathids is not available. As a result it is of very limited stratigraphical use.
The out-dated studies of Taylor and the too brief publications of the Bedfords, have
not allowed specialists in other countries to make precise comparisons and possible
correlations, between their material and that of Australia. An attempt was made on
the limited material available in the Ting collection (F. & M. Debrenne 1960). It
was concluded, from the majority of genera appearing at Beltana, that the fauna
belonged to the Aldanian and Lower Lenian stages. Since then, I have become
acquainted with Russian work that amends the stratigraphical correlation between
the Southern Siberia geosyncline and the Siberian platform, and gives detailed lists of
revised Archaeocyatha faunas from level to level.
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 369
These data, together with the present study, confirm that the Ajax Limestone is
situated in the lower half of the Lower Cambrian ; to be precise, the fauna is similar to
that of the Kameshki horizon and also has a certain affinity with those forms found in
the succeeding Sanashtygkol horizon. This opinion is based on the presence, in the
Ajax fauna, of forms having advanced evolutionary stages of outer and inner walls,
or intervallar structures, together with other simple forms known from the Bazaikha
horizon, such as Monocyathus, Alphacyathus, Ajacicyathus, Robustocyathus and
Erismacoscinus .
As examples of the advanced forms, one can show the presence of genera with :
1. Outer walls with simple tumuli (Tumuliolynthus , Ethmocoscinus] , or multi-
perforate (A lataucyathus] .
According to Rozanov (1963 : 8) genera with tumuli mainly arise in the
Kameshki horizon.
2. Double porous outer wall (Tomocyathus, Polycoscinus}.
This feature appears at the end of the Bazaikha horizon and genera that
possess it increase in numbers through the Kameshki and Sanashtygkol horizons.
3. Inner wall with complex annular rings.
This is seen in Cyathocricus annulispinosus (Vologdin) found in the Kameshki
horizon of Eastern Sajan, and the Australian species annulatus (Bedford)
regarded as belonging to Salairocyathus, which occurs in the Sanashtygkol
horizon.
4. Inner wall with branching pore-tubes (Zonacyathus).
Three Russian species probably belong to this genus ; poletaevae, vermiculatum
and flexum. They and the corresponding colonial genus Sajanocyathus are of
Sanashtygkol age.
5. Radial septa with few, or no pores (Cyathocricus, Ethmocyathus, Zonacyathus,
Robustocyathus subacutus and Archaeocyathellus (Stapicyathus] stapipora}.
Zhuravleva (1960 : 147) and Rozanov & Missarzhevsky (1966 : 71) consider
that the decrease or loss of septal porosity is an evolutionary character of the
Regularia. The pores of the plates are often different to the septal pores in
Australian Coscinocyathus, whereas in specimens found at lower levels in the
Cambrian (e.g. Soussian horizon in Morrocco) they are very similar. The first
appearance of any difference between the two occurs in the Sardinian limestones,
contemporaneous with the Botoma Stage of Southern Siberia.
LOWER CAMBRIAN
These correlations are taken from Table II of Walter (1967), who used information
given in a personal communication (Sept. 1966) to M. F. Glaessner by A. Yu. Rozanov.
SIBERIAN PLATFORM GEOSYNCLINE OF SOUTHERN SIBERIA
Lower subdivision
f Sunnagin fKundatsk horizon
TOMMOT STAGE < T, I ^ ., , , .
\Kenyada ALDAN STAGED Bazaikha horizon
Atdaban (^ Kameshki horizon
Tarynsk
Sinsko-Kutorgina BOTOMA STAGE Sanashtygkol horizon
370 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
Upper subdivision
Ketema LENA STAGE Solontsov horizon
Elanka Obruchev horizon
Similar studies of evolutionary stages, have not been made for the Irregularia, but
it is possible to make the following points.
1. The first Irregularia have scattered skeletal elements, rods, or plates, as in
Dictyocyathus, or Protopharetra and are of Soussian age. They then follow two
particular paths.
a. The plates of Protopharetra gave rise to the crude pseudo-septa of Volva-
cyathus-iype, where the pores occupy a greater surface area than the
skeletal tissue; then into the poorly developed pseudosepta shown in the
first Metaldetes of Sardinia ; followed by the definite radial septa of the true
Metaldetes of Australia and ending with the Cambrocyathus-type, which
occurs in the upper half of the Lower Cambrian.
b. The rods of Dictyocyathus, on the other hand, form the regular scaffolding
of Flindersicyathus and Pycnoidocyathus types, which are comparable to the
intervallar structure of Tabellaecyathidae Fonin, characteristic of the
Sanashtygkol horizon.
2. Amongst the Ajax fossils one finds Syringocnema, a genus characterized by
prismatic loculi in the intervallum. In the USSR the species referred to this
genus are limited to the Sanashtygkol horizon.
3. The walls of the Irregularia are not so diverse as those of the Regularia. In the
older forms of Dictyocyathus and Protopharetra they are not independent of the
intervaller structural elements. This is attained in the genus Flindersicyathus,
with its simply porous outer wall and inner wall having pore-tubes, the lower
parts of which are modified and stretched. The double porous inner wall is seen
in the genera Metaldetes, Metafungia and Metacoscinus. Such structures indi-
cate the development of more advanced features, e.g. lamellae and tabulae, that
are known in the Sanashtygkol horizon (Fonin 1963).
All these considerations show that there are clear affinities between the Ajax faunas
and those of the Kameshki and Sanashtygkol horizons. The absence of forms with
clathrate outer walls, or inner walls of true Ethmophyllum-type, shows that the
Sanashtygkol fauna was not well established in Australia at the time of the Ajax
limestone. The fauna is, therefore, probably of Upper Kameshki and Lower
Sanashtygkol age, that is to say the middle part of the lower division of the Lower
Cambrian.
Since this conclusion was first written Walter (1967), in his study on the usefulness
of Archaeocyatha for zoning and correlation in the Lower Cambrian of the Adelaide
Geosyncline, has correlated the various faunas in the Hawker Group of South
Australia with those of Siberia. Using genera as the basis of such correlation, he has
assigned a Sanashtygkol age to the Oraparinna Shale; the latter occurs above the
LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA 371
Hawker Group
Wilpena Group
S\\\\\'
Wirrealpa Lst.
Billy Creek Form'n.
Marina Graywacke
Oraparinna Shale
Bunkers Sdst.
Parara Lst.
Wilkawillina Lst.
Parachilna Form'n.
Pound Qtzte.
Ajax Lst,
FIG. 15. Lower Cambrian Sediments, Flinders Ranges; S. Australia.
After Dalgarno (1964) & Walter (1967).
Wilkawillina Limestone of which, in the Mt. Scott Range area, the Ajax Limestone is
said to be the equivalent (Dalgarno 1964 : 136). Commenting on the Ajax Limestone
Walter (p. 146) suggests that the presence of Syringocnema, Flindersicyathus and
Pycnoidocyathus indicate correlation with the upper part of the Hawker Group in the
Wilkawillina Gorge section and concludes that, in the Ajax Mine area, the Ajax Lime-
stone may be another bioherm. This evidence would appear to support my con-
clusions as to the age of the fauna of the Ajax Limestone.
IV. SUMMARY
The material now in the British Museum (Natural History), collected and
originally described by R. & W. R. Bedford, has been revised according to current
classification.
372 LOWER CAMBRIAN ARCH AEOC Y ATH A FROM AUSTRALIA
It is considered necessary to create a separate family within the Ajacicyathacea for
the genus Ethmocyathus, which is characterized by a secondary wall of annular plates
on the central cavity side of the inner wall. A new genus, Cyathocricus, is introduced
for Bronchocyathus Bedford, which did not conform to the rules of zoological nomen-
clature, and includes those forms belonging to the Bronchocyathidae that have a more
complex inner wall than Cyclocyathus. Two species are assigned to the new genus.
A new subgenus Loculicyathus (Loculicyathellus] is distinguished from Loculicyathus
(Loculicyathus} by the longitudinal corrugations occurring between the septa on the
outer wall. Two new subgenera are introduced for the smooth inner wall forms of
Tomocyathus: T. (Erugatocyathus) and Alataucyathus : A. (Anaptyctocyathus}. A
new species of the latter is erected, A. (A.} flabellus, which is distinguished from A.
(A.} cribripora by its bowl-shaped cup, dimensions and arrangement of wall pores.
In the broader classification, the family Acanthinocyathidae is distinguished from
the Dokidocyathidae, and the family Bronchocyathidae is re-introduced to take the
new genus Cyathocricus. The genus Flindersicyathus is included in the re-introduced
family Flindersicyathidae, as there are many characters of this genus which distin-
guish it from Archaeocyathus. Pycnoidocyathus and Flindersicyathus are treated as
subgenera until the Ajax Mine material in both Princeton and Adelaide is re-studied.
Owing to both the poor preservation and limited number of some specimens, it has
been impossible to see sufficient structural details for exact determination and the
true position of several genera remains uncertain. However, the good condition of
other material has enabled a thorough study of the internal skeleton to be made, with
the result that new types of structure have been discovered, e.g. the annular form of
the inner wall in Salairocyathus (S.) annulatus.
From the stratigraphical evidence, it is concluded that the fauna is of middle Lower
Cambrian age, i.e. upper Kameshki — lower Sanashtygkol horizons, and this has been
borne out by subsequent work.
V. ACKNOWLEDGMENTS
I wish to thank Dr. E. I. White and Dr. H. W. Ball for the opportunity of studying
the Bedford collection in the Department of Palaeontology. I would also like to
thank Dr. Ball and Mr. S. Ware, for the generous help and information given to me
whilst examining this collection. I am also extremely grateful to Professor M. F.
Glaessner and his colleagues of the University of Adelaide and at the Geological
Survey of South Australia, for allowing me to reproduce maps and stratigraphical
information in text-figs. 13, 14 and 15. In addition, thanks are also due to my
husband M. M. Debrenne and Mr. P. Green of the Photographic Studio at the B.M.
(N.H.) for the photographs. Finally, I wish to thank Dr. Dorothy Hill, of the Uni-
versity of Queensland, for her encouragement and advice, and for permission to
reproduce some of the photographs of Bedford material included in her monograph
on Antarctic archaeocyathids.
LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA 373
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— — 1932. The Archaeocyathinae of Siberia. 2. Fossils of the Cambrian Limestones of the
Altai Mountains. Izv. geol.-razved. Ob' 'ed., Moscow. 1932 : 106, 14 pis., 46 figs. [In
Russian with English translation.]
- 1937. Arkheotsiaty i vodorosli yuzhnogo sklono anaborskogo massiva. Trudy arkt.
nauchno-issled. Inst., Leningrad. 91 : 9-46. [French translation S.I.G. 1434.]
- 1939. Middle Cambrian Archaeocyatha and Algae from the South Urals. Problem-
Paleont., Moscow. [In Russian with English translation.] 5 : 209-276, 12 pis., 12 figs.
— 1940. Les archeocyathes et les algues des calcaires cambriens de la Mongolie et de la Tuva.
Trudy mongol' Kom., 34, 10 : 268. [In Russian with English summary.]
- i94oa. (Ed.) i. Porifera, Archeocyatha. In " Atlas of the leading forms of the fossil
faunas of the USSR i. Cambrian. " Trudy vses. nauchno-issled. geol. Inst., Moscow.
1940 : 23-97, 31 pis., 85 figs. [In Russian with French transl. SIG 1446.]
- 1956. Classification du type Archaeocyatha. Dokl. Akad. Nauk SSSR. Moscow. Ill,
4 : 877-880. (In Russian, with French translation SIG No. 1510.)
- 1957. Les Archaeocyatha et leur signification stratigraphique. Annls. Cent. £tud.
Docum. paleont., Paris. 23, 2 : 34-73, 22 pis. [French translation of Acta palaeontologica
sinica, 5, 2 : 173-222.]
VOLOGDIN, A. G. 1961. Arkheotsiaty i ikh stratigraficheskoe znachenie. In " El sistema
Cambrico, su paleogeografia y el problema de su base. Part 3. Asia. " XXth Int. Geol.
Congr. Mexico, 1956 : 173-199.
VOLOGDIN, A. G. 1962. [Archaeocyatha and algae of the Cambrian in the Baikal Highlands.]
Trudy paleont. Inst., Moscow 93 : 3-116, 21 pis., 21 figs. [In Russian.]
— - — ig62a. Tip Archaeocyatha, Arkheotsiaty. In ORLOV, Y. A. (ed. " Osnovy Paleontologii. "
2 [Spongia, Archaeocyatha, Coelenterata, Vermes]. Akad. Nauk USSR, Moscow, 1962:
89-139. [In Russian.]
--- ig62b. [The anatomy of the Archaeocyathids.] Paleont. Zh., Moscow. 1962, 2 : 9-20.
[In Russian with English translation in Int. Geol. Rev., Washington. 5, 12 : 1635-1647.]
376 LOWER CAMBRIAN ARCH AEOCY ATH A FROM AUSTRALIA
VOLOGDIN, A. G. 1963. [Late Middle Cambrian Archaeocyathids from the Amga River basin (on the
Siberian Platform)]. Dokl. Akad. Nauk SSSR, 151, 4 : 946-949. [In Russian, with English
translation in Dokl. (Proc.) Acad. Sci. URSS, Earth Sci., Washington. 151, 1963 : 199-202.]
WALTER, M. R. 1967. Archaeocyatha and the biostratigraphy of the Lower Cambrian Hawker
Group, South Australia. /. geol. Soc. Aust., 14, i : 139-152, 2 pis.
YAKOVLEV, V. N. 1956. [On some poorly expressed characters of the structure of Archaeo-
lynthus Taylor and its possible genetic connection with Echinodermata.] Dokl. Akad. Nauk
SSSR, Moscow. 109, 4 : 855-857, i pi. [In Russian.]
ZHURAVLEVA, I. T. 1949. Certaines donn6es sur la structure de calice chez les representants
du genre Rhabdocyathus Toll. Dokl. Akad. Nauk SSSR, Moscow. 67, 3,: 547-550, 2 figs.
[In Russian. French translation SIG 240.]
- 1951. Sur 1' individuality des calices brises d'Archeocyathes Reguliers et de « larves »
d'Archeocyathes. Dokl. Akad. Nauk SSSR, Moscow. 80, i : 97-100, 3 figs. [In Russian,
French translation SIG No. 407.]
— 1955. Contribution a la connaissance des Arch6ocyathes de Siberie. Dokl. Akad. Nauk.
SSSR, Moscow. 104, 4 : 626-629, i pi-, i fig. [In Russian, French translation SIG 1346.]
1960. Archaeocyathi of the Siberian Platform. Izd. Akad. Nauk SSSR, Inst. Geol. Geofiz.,
Moscow. 1960 : 346, 33 pis., t.-figs. [In Russian.]
— i96oa See under KHALFIN, L. L. (Ed.). 1960.
— 1961. See under Musatov, D. I. et al., 1961.
- 1963. Archaeocyatha of Siberia: single-walled archaeocyatha. Orders Monocyathida —
Rhizacyathida. Izd. Akad. Nauk SSSR, Inst. Geol. Geofiz., Moscow. 1963 : 139, 87 pis.
[In Russian.]
- 1964. See under REPINA, L. N. (Ed.). 1964 : 166-251.
1 964 a. See Below.
ZHURAVLEVA, I. T., KONYUSHOV, K. N. & ROZANOV, A. Yu. 1964. Archaeocyatha of Siberia:
The two-walled Archaeocyatha. Izd. Akad. Nauk SSR, Inst. Geol. Geofiz., Moscow.
1964 : 1 66, 16 pis., 75 figs. [In Russian.]
APPENDIX
GLOSSARY
APEX initial part of cup.
CENTRAL CAVITY space inside the inner wall.
DISSEPIMENTS non-skeletal vesicular connections between the septa.
ENDOTHECA tissue formed on the central cavity side of the inner wall.
EXOTHECA diverse external outgrowths from the intervallum and outer wall.
INTERSEPT/INTERSEPTUM space between two adjacent septa.
INTERTABULUM space between 2 successive tabulae.
INTERVALLUM space between the inner and outer walls.
LINTEAUX skeletal elements around the pores.
LOCULUS space limited by the two walls, two adjacent septa and on occasion
by two successive tabulae.
PELLIS thin calcareous sheath, porous or not, outside the walls of some
genera.
PORES simple perforations in the skeletal elements.
SEPTA porous, radial vertical plates connecting the inner and outer walls.
SYNAPTICULAE horizontal rods between 2 neighbouring septa.
TABULAE straight, or curved; porous, or pectinate plates that cross the
intervallum.
TAENIAE thick, skeletal; irregular, or waved; sometimes gondola-shaped,
plates, that radially subdivide the intervallum and are only found
in the Irregularia.
TUMULI wall protuberances.
EXPLANATION OF PLATES
All the figured material is in the collections of the British Museum (Natural
History).
PLATE i
FIG. i. Tumuliolynthus irregularis (R. & W. R. Bedford). Holotype 84141. Outer
view of the wall and tumuli. X 15.
FIG. 2. Alphacyathus cf. annularis R. & W. R. Bedford. S 4822. Etched longitudinal
section showing the outer wall, the inner wall and the horizontal rods. X 15.
FIG. 3. Monocyathus porosus R. & W. R. Bedford. Lectotype S 4140. Cup showing
outer view of the wall. X 6.
FIG. 4. Monocyathus mellifer R. & W. R. Bedford. 84821. Outer view of the wall.
X 6.
FIG. 5. Archaeocyathellus (Stapicyathus) stapipora (Taylor). 84733. Detail of
inner wall. X 25.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE i
GEOL. 17, 7
PLATE 2
Acanthinocyathus apertus R. & W. R. Bedford
FIG. i. Paratype S 4167. View of part of outer wall showing pores and position of spines.
X 10 approx.
FIG. 2. Paratype S 4167. View of part of outer wall showing lateral view of a spine.
X 6 approx.
FIG. 3. Holotype S 4166. Another view of outer wall showing the occurrence of the spines.
X 10 approx.
FIG. 4. Paratype S 4168. View of inner wall. X 4.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 2
GEOL. 17, 7
PLATE 3
Loculicyathus (Loculicyathellus) floreus (R. & W. R. Bedford)
Holotype S 4144.
FIG. i. Inner wall and inner part of septa, x 10.
FIG. 2. Outer wall on two intersepts. x 10.
FIG. 4. Upper transverse section, x 10.
Ethmocoscinus papillipora (R. & W. R. Bedford)
Holotype S 4164.
FIG. 3. Part of transverse section, x 15.
Robustocyathus tnagnipora (R. & W. R. Bedford)
Holotype S 4146.
FIG. 5. Inner wall, septa and outer wall seen from the central cavity, x 5.
Robustocyathus subacutus (R. & W. R. Bedford)
Holotype S 4142.
FIG. 6. Transverse section, x 5.
FIG. 7. View from central cavity of inner wall, septa, x 5.
Bull. Br. Mus. nat. Hist. (GeoU 17, 7
PLATE 3
PLATE 4
Zonacyathus retezona (Taylor)
FIG. I. 84731. Detail of inner wall, x 12-5.
FIG. 2. S 4764. Oblique transverse section, x 6.
FIG. 3. S 4353. Etched longitudinal section showing inner wall, septa and outer wall, x 6.
Zonacyathus retevallum (R. & W. R. Bedford)
FIG. 4. Holotype S 4147. Detail of inner wall. X 15.
FIG. 5. 84726. Transverse section, x 6.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
1
PLATE 4
PLATE 5
Ethmocyathus lineatus R. & W. R. Bedford
Holotype S 4149.
FIG. i. Detail of inner wall showing lozenge pores and tabellae. x 10.
FIG. 2. Porosity of septa, x 25.
FIG. 3. Inner view of outer wall. X4O.
Cyathocricus tracheodentatus R. & W. R. Bedford
Holotype S 4148.
FIG. 4. Lower transverse section, x 6.
FIG. 5. Etched longitudinal section showing the inner wall, x 6.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
1
PLATE 5
4?'^:
jf ^ ' ^ *^*^ "•*»
^T*~2*I-*-
^V*"?!*&,.
fsir
^p.
PLATE 6
Cyathocricus dentatus (Taylor)
FIG. i. S 4753. Longitudinal section showing the rings and teeth (note that the septa are
only porous near the outer wall) and outer wall, x 30.
FIG. 2. S 4753. Etched transverse section, x 10.
FIG. 3. S 4355. Etched longitudinal section showing the outer wall, the septa and the inner
wall (at the upper part of the figure), x 8.
Cyathocricus tracheodentatus (R. & W. R. Bedford)
FIG. 4. S 4754. Detail of inner wall, x 25.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 6
PLATE 7
Erismacoscinus equivallum (Taylor)
Holotype S 198.
FIG. i. Detail of one tabula [on opposite side to Fig. 2]. X 15.
FIG. 2. Transverse section showing inner wall pores and tabula, x 3.
FIG. 3. Detail of outer wall, x 8.
FIG. 5. Detail of inner wall. X 15.
Erismacoscinus quadratus (R. & W. R. Bedford)
Holotype S 4157.
FIG. 4. Etched oblique longitudinal section, x 5.
Bull. Br. Mus. not. Hist. (Geol.) 17, 7
PLATE 7
PLATE 8
Erismacoscinus cellularis (R. & W. R. Bedford)
Holotype S 4162.
FIG. i. Detail of septa and inner wall (on right). X 15.
FIG. 4. Oblique transverse section, x 6.
Erismacoscinus rugosus (R. & W. R. Bedford)
Holotype S 4152.
FIG. 2. Outer view of outer wall, x 10.
FIG. 3. Oblique longitudinal section, x 10.
Erismacoscinus quadratus (R. & W. R. Bedford)
FIG. 5. Holotype S 4157. Transverse section and detail of the inner wall, x 10.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
1
PLATE 8
PLATE 9
Erismacoscinus textilis (R. & W. R. Bedford)
Paratype S 4156.
FIG. I. Detail of outer wall (lower right), septa, tabulae, vesicular tissue (centre) and inner
wall (above left). x6.
FIG. 3. Bowl shape. X 1-5.
Holotype S 4155.
FIG. 2. Cylindrical shape, x 4.
FIG. 4. Transverse section, x 5.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 9
PLATE 10
Alataucyathus (Anaptyctocyathus) flabellus sp. nov.
Holotype S 4161.
FIG. i. Outer view. xa.
Eristnacoscinus retifer (R. & W. R. Bedford)
Holotype S 4159.
FIG. 2. Etched inner wall seen from central cavity, x 10.
FIG. 3. Transverse section, x 5.
Eristnacoscinus petersi (R. & W. R. Bedford)
Holotype S 4158.
FIG. 4. Transverse section, x 10.
FIG. 5. Longitudinal section, x 10.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 10
PLATE ii
Alataucyathus (Anaptyctocyathus) cribripora (R. & W. R. Bedford)
Holotype S 4160.
FIG. i. Detail of outer wall, external view of pores and micropores. X25.
FIG. 3. Longitudinal section, x 10.
Tomocyathus (Erugatocyathus) papillatus (R. & W. R. Bedford)
Holotype S 4153.
FIG. 2. Upper transverse section. X5-
Salairocyathus (Salairocyathus) annulatus (R. & W. R. Bedford)
Holotype S 4163.
FIG. 4. Detailed longitudinal section showing the inner wall, x 5.
Bull. Br. Mus. not. Hist. (Geol.) 17, 7
PLATE ii
GEOL. 17, 7
PLATE 12
Pinacocyathus spicularis R. & W. R. Bedford
Holotype S 4169
FIG. i. Transverse section, x 7.
FIG. 2. Outer view showing the scaffolding of rods, x 7.
Flindersicyathus (Flindersicyathus) graphicus (R. & W. R. Bedford)
FIG. 3. Paratype 84171. Longitudinal section through an unwaved taenia (with central
cavity to the left), x 6.
FIG. 4. Paratype 84171. Longitudinal section showing an inner view of the inner wall, x 6.
FIG. 5. Holotype S 4170. Transverse section, x 6.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 12
PLATE 13
Metaldetes taylori (R. & W. R. Bedford)
Holotype S 4185-4187 (three pieces of the same specimen)
FIG. i. S 4185. Lower part; external view. X2-2 approx.
FIG. 2. S 4186. Upper part; transverse section, x 1-8.
FIG. 3. S 4186. Upper part; oblique and longitudinal sections with the septa running N-S.
X 2 approx.
These photographs are used by courtesy of the British Museum (Nat. Hist.) and Dr. Dorothy
Hill.
tCoscinocyathus unilinearis (R. & W. R. Bedford)
FIG. 4. Holotype S 4165. View of both inner and outer walls and intervallum. x 5.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 13
f^ • x»'. j'if&itf^f^*) , * * ''' ' * i * ^^*^i*» '*
T*!'. ', * • v ^x2v*4 '&*£* ^>*^-t^**> x »
'%.'.**» ' * <-./,<• ^-^.^^^v*'*^^*.**. «•
PLATE 14
Flindersicyathus (F.) speciosus (R. & W. R. Bedford)
Holotype S 4175.
FIG. i. Lower transverse section. X5-
FIG. 3. Etched longitudinal section showing inner wall tubes, x 5.
Flindersicyathus (F.) major (R. & W. R. Bedford)
FIG. 2. Holotype S 4174. Etched longitudinal section showing the bending of flat taeniae
(on right) and the widened tubes of the inner wall (on left), x 6.
Flindersicyathus (F.) irregularis (Taylor)
FIG. 4. S 4763. Part of transverse section showing thickened inner side of taenia. X 10.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 14
PLATE 15
FIG. i. Flindersicyathus (Pycnoidocyathus) synapticulosus (Taylor) 8208. Part of
transverse section. X 2-5.
FIG. 2. Flindersicyathus (P.) simplex (Taylor). Paratype 84824. Part of longitudinal
and transverse sections. X 3-
FIG. 3. Flindersicyathus (P.) maximipora (R. & W. R. Bedford). Holotype S 4150.
Part of an etched longitudinal section showing extended pore-tubes of the inner wall and unwaved
taeniae. X 3-
FIG 4. Flindersicyathus (P.) vicinisepta (R. & W. R. Bedford). S 4825. Part of longi-
tudinal section showing the inner wall. X 5.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 15
PLATE 16
Mctaldetes dissepimentalis (Taylor)
FIG. i. 84176. Lateral view of non- waved taeniae. x6.
FIG. 2. 84182. Part of transverse section. x6.
FIG. 3. 8 4176. Outer view of outer wall. X 6.
Metaldetes irregularis (R. & W. R. Bedford)
FIG. 4. Paratype S 4190. Outer view of outer wall, x 6.
Bull. Br. Mus. not. Hist. (Geol.) 17, 7
PLATE 16
PLATE 17
Metacoscinus reteseptatus R. & W. R. Bedford
FIG. i. S 4194. Outer view of outer wall, x 6.
FIG. 2. S 4191. Part of transverse section showing a tabula, x 10.
FIG. 3. S 4195. Inner wall seen from the central cavity. x6.
FIG. 4. S 4194. Section along a taenia (central cavity on left). x6.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 17
PLATE 18
Metafungia reticulata R. & W. R. Bedford
Holotype S 4184.
FIG. i. Inner wall of cup (on left). X4-
FIG. 2. Outer wall, x 6.
FIG. 3. Transverse section. X3.
Photographs by courtesy of the British Museum (Nat. Hist.) and Dr. Dorothy Hill.
Pycnoidocoscinus pycnoideum R. & W. R. Bedford
S 4832.
FIG. 4. Part of transverse section showing thickened taeniae. X5.
FIG. 5. Longitudinal section showing inner wall, x 5.
FIG. 6. Section showing area of fig. 7. X5-
FIG. 7. Detail of tabulae pores, x 30.
FIG. 8 . Detail of taeniae and tabulae, x 30.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 7
PLATE 18
PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING
SOME BATHONIAN OSTRACODA
OF ENGLAND WITH A REVISION
OF THE JONES 1884 AND JONES
& SHERBORN 1888 COLLECTIONS
R. H. BATE
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 8
LONDON: 1969
SOME BATHONIAN OSTRACODA OF ENGLAND
WITH A REVISION OF THE JONES 1884 AND
JONES & SHERBORN 1888 COLLECTIONS
BY
RAYMOND HOLMES BATE
Pp. 377-437; 16 plates, 18 text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 17 No. 8
LONDON: 1969
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 17, No. 8 of the Geological
(Palaeontological] series. The abbreviated titles of
periodicals cited follow those of the World List of
Scientific Periodicals.
World List abbreviation
Bull. Br. Mus. nat. Hist. (Geol.).
Trustees of the British Museum (Natural History) 1969
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 16 May, 1969 Price £3 55.
SOME BATHONIAN OSTRACODA OF ENGLAND
WITH A REVISION OF THE JONES 1884 AND
JONES & SHERBORN 1888 COLLECTIONS
By R. H. BATE
CONTENTS
Page
I. INTRODUCTION ......... 379
II. THE T. R. JONES COLLECTION ...... 380
III. THE T. R. JONES & C. D. SHERBORN AND THE H. H. WINWOOD
COLLECTIONS ......... 394
IV. THE J. F. BLAKE COLLECTION ...... 427
V. REFERENCES .......... 435
SYNOPSIS
Three Bathonian ostracod Collections from the Fuller's Earth Clay of Bath, Somerset, and
one of similar age (the T. R. Jones Collection, published in 1884) from the Richmond borehole,
Surrey, are described and illustrated, that of J. F. Blake for the first time. The H. H. Win wood
Collection is identified as syntypic material of the T. R. Jones & C. D. Sherborn Collection,
published in 1888. Sixty- three ostracod species are described of which six are new, and six
are left under open nomenclature. Nineteen species are listed as lost. Two genera, Hadro-
cytheridea and Hekistocy there, are new.
I. INTRODUCTION
IN 1884 T. R. Jones described the foraminifera and Ostracoda from a deep boring at
Richmond, Surrey. The ostracods were described from three levels and, apart from
Pontocyprella harrisiana and Schuleridea jonesiana which are Cretaceous contami-
nants of the collection, are Bathonian in age.
Four years later, Jones & Sherborn (1888) described a much larger ostracod fauna
from the Bathonian of Midford, near Bath. The samples of Blue and Yellow Fuller's
Earth Clay were given to Jones and Sherborn by the Rev. H. H. Win wood (see Jones
& Sherborn 1886).
The collections of Jones 1884, and of Jones & Sherborn 1888, are of considerable
importance for present-day work. Because of this it is proposed here to re-describe
and re-illustrate the species and to designate types where necessary; a few species
originally described by Jones & Sherborn are missing from the collections and are
presumed lost.
Two additional collections from the Fuller's Earth of Midford, near Bath, are
included because they add to the faunal list and in the case of the Blake collection
record the presence, for the first time in England, of a number of species previously
only described from N.W. France (Oertli 1959). The most important of the collec-
tions, however, is that presented by the Rev. H. H. Winwood. A label stating that
GEOL. 17, 8 23
380 REVISION OF SOME
these specimens are duplicates of the originals figured in 1888 was left with the collec-
tion by C. D. Sherborn. As the specimens figured by Jones & Sherborn were ex-
tracted from material given them by Winwood, there would appear to be no doubt
that this collection represents syntypic material.
The Fuller's Earth Clay of Midford, near Bath, was placed by Arkell (1956 : 28)
in the Oppelia aspidoides Zone. Torrens (1967 : 83), however, assigned it to the zone
of Prohecticoceras retrocostatum, basal Upper Bathonian and below the aspidoides
Zone.
The terminology and classification employed throughout this paper are those of
Moore (1961) and Bate (1963). All the material described is in the Department of
Palaeontology, British Museum (Natural History).
II. THE T. R. JONES COLLECTION
Order PODOCOPIDA Miiller 1894
Suborder PLATYCOPINA Sars 1866
Family CYTHERELLIDAE Sars 1866
Genus CYTHERELLA Jones 1849
Cytherella symmetrica Jones
(PI. I, figS. I, 2)
1884 Cytherella symmetrica Jones : 768, pi. 34, fig. 42.
1884 Cytherella subovata Jones : 773, pi. 34, fig. 43.
DIAGNOSIS. Cytherella with elongate-oval carapace. Left valve uniformly over-
reached by larger right valve. Greatest height of carapace in posterior third. Shell
surface very finely punctate. Dimorphic: Adult female of length 0-85 mm., male
of length 0-90 mm.
LECTOTYPE. IN.43503, female right valve; Richmond boring, depth 1,151 ft. 6 ins.
PARALECTOTYPES. IN.43496, male left valve, and 1.941, female carapace; depth
1,205 ft.
DESCRIPTION. Both left and right valves are elongate-oval in outline with well-
rounded anterior and posterior margins. Ventral margin straight to slightly convex,
dorsal margin strongly convex in the posterior third (region of greatest height) but
straightens out anteriorly. Greatest length through mid-point. Right valve
larger than the left which it overreaches evenly all-round. Shell surface very finely
punctate, though the state of preservation does not always show this. Selvage in
the left valve strongly developed to form a ridge which fits into a corresponding
groove in the right.
DIMENSIONS. IN.43503. Female right valve and lectotype of Cytherella sym-
metrica Jones, length 0-85 mm. ; height 0-52 mm. 11^.43496. Male left valve and
ENGLISH BATHONIAN OSTRACODA 381
lectotype of Cytherella subovata Jones, length 0-90 mm. ; height 0-50 mm. 1.941.
Female carapace, length 0-77 mm. ; height 0-49 mm. ; width 0-36 mm.
REMARKS. The figured specimens of C. symmetrical and of C. subovata are missing
from the Jones Collection and are presumed lost. Unfigured syntypes have therefore
been selected as lectotypes. C. subovata has been recognized as the male dimorph
and C. symmetrica as the female dimorph of the species Cytherella symmetrica.
Oval species of Cytherella look very much alike, and C. symmetrica is very similar
to C. suprajurassica Oertli (1957 : 649, pi. i, figs. i-io). The latter, however, is a
smaller species with a greater mid-dorsal projection of the right valve over the left.
In C. symmetrica the overreach of the right valve is much more uniformly developed
along the entire dorsal margin.
Genus CYTHERELLOIDEA Alexander 1929
Cytherelloidea jugosa (Jones)
(PI. i, figs. 3, 4; Text-fig, i)
1884 Cytherella jugosa Jones : 773, pi. 34, fig. 44.
DIAGNOSIS. Carapace sub-rectangular in outline with broadly rounded anterior
and posterior margins. Ventral margin strongly incurved. Ornamentation consists
of a broad peripheral ridge and a thick, central sigmoid ridge. Shell surface finely
punctate.
LECTOTYPE. ^.43497, female right valve (figured Jones 1884) ; Richmond boring,
1,205 ft.
PARALECTOTYPE. 1.2311, female carapace; Richmond boring, depth not recorded.
DESCRIPTION. Carapace sub-rectangular in outline with well rounded anterior
and posterior margins. Dorsal margin broadly convex with a slight concavity just
anterior of mid-point. Ventral margin strongly incurved. Greatest length through
mid-point ; greatest height median, despite the strong concavity of the ventral margin.
Greatest width at the posterior margin. The two specimens available are female
dimorphs possessing two posterior swellings, of which the lowermost is the more
prominent. Shell surface finely punctate, further ornamented by a broad peripheral
ridge extending completely around each valve and enclosing a sigmoid inner ridge
which commences at the anterior inner edge of the peripheral ridge, bends upwards,
then curves down below the mid-dorsal muscle scar depression, finally curving
upwards again to die out before reaching the peripheral ridge in the region ol the
postero-dorsal angle. Right valve larger than the left, which it overreaches anteriorly
and overlaps ventrally and dorsally, especially anterodorsally where there is a slight
concavity in the margin. Only internal details of the right valve observed: Muscle
scars situated on a slightly raised dorso-median boss, are typical of the genus; a
groove extends around the posterior and along the dorsal margin for the reception
of the left valve. Two posterior cavities correspond to the swellings seen on the
outside of the valve.
382 REVISION OF SOME
DIMENSIONS.
IN-43497. Female right valve, length 0-55 mm. ; height 0-30 mm.
1.2311. Female carapace, length 0-56 mm. ; height 0-34 mm. ; width 0-23 mm.
REMARKS. C. jugosa is similar to C. paraweberi Oertli (1957 : 651, pi. i, figs.
12-15) but differs in being rounded rather than angular postero-dorsally, in being
more noticeably constricted mid-dorsally, and in being narrower anteriorly where
only the right valve tends to project. Laterally, the ribbing of C. jugosa is very
much broader. A weak reticulation may be observed in C. paraweberi on the lateral
surface.
FIG. i. Cytherelloidea jugosa (Jones). Female right valve. Lectotype 11^.43497. X 135.
FIG. 2. Cytherelloidea paraweberi Oertli. Right side, female carapace, 10.3527. x 130.
Stratigraphically, C. jugosa predates C. weberi Steghaus (1951 : 207, pi. 14, figs.
3-6) and C. paraweberi Oertli, both of which appear to be restricted to the
Kimmeridgian. The close similarity in ornamentation between these species suggests
a phylogenetic relationship (Text-figs, i, 2). This is probably also true for the
Bajocian species., Cytherelloidea eastfieldensis Bate (19630 : 25, pi. i, figs. 1-5, Text-
fig, i) which is easily distinguishable by the mid-dorsal discontinuity of the peripheral
ridge.
ENGLISH BATHONIAN OSTRACODA 383
Suborder PODOCOPINA Sars 1866
Superfamily BAIRDIAGEA Sars 1888
Family BAIRDIIDAE Sars 1888
Genus BAIRDIA McCoy 1844
Bairdia hilda Jones
(PL i, figs. 5, 6; PL 4, fig. 5)
1884 Bairdia hilda Jones : 771, pi. 34, fig. 20.
1888 Bairdia fullonica Jones & Sherborn : 253, pi. 5, fig. ^a-c.
1948 Bairdia cf. hilda Jones; Sylvester-Bradley : 199, Text-fig. 5.
1963 Bairdia hilda Jones; Bate : 188, pi. 2, figs. 9-12, pi. 3, figs. 1-4.
1964 Bairdia hilda Jones; Bate : 8.
DIAGNOSIS. Carapace subdeltoid laterally, strongly convex dorsally. Dorsal
margin high with steeply inclined anterior and posterior slopes. Posterior end
acuminate, upturned. Shell surface finely punctate.
LECTOTYPE. IN.4I95I, single left valve figured Jones (1884) Richmond boring,
depth 1,205 ft.
PARALECTOTYPES. 10.3608-16 and 10.3620-6, carapaces and single valves from
a depth of 1,205 ft.
REMARKS. Full descriptions of this species have been given previously by
Sylvester-Bradley (1948) and Bate (1963). However, as the measurements of the
types were not included these are given here.
DIMENSIONS. IN.4I95I, left valve, length 0-93 mm. ; height 0-48 mm. 10.3608.
carapace, length 0-77 mm. ; height 0-48 mm. ; width 0-34 mm. 10.3610, carapace,
length o-qo mm. ; height 0-47 mm. ; width 0-34 mm.
Bairdia juddiana Jones
(PL i, fig. 7)
1884 Bairdia juddiana Jones : 767, pi. 34, fig. 18.
DIAGNOSIS. Carapace globose, convex in dorsal view. Shell surface finely
punctate. Antero-dorsal and postero-dorsal slopes convex. Left valve distinctly
overlaps the right along the dorsal and ventral margins, particularly mid-ventrally.
Caudal process slightly upturned. Ventral margin angled with flattened median
portion and steeply sloping antero-ventral and postero- ventral slopes.
HOLOTYPE. IN.435o6, carapace, figured Jones (1884), Richmond boring, depth
1,151 ft.-i,i5i ft. 6 in.
DESCRIPTION. Carapace rather globose in lateral view with convex antero-dorsal
and postero-dorsal slopes. Anterior end rounded, posterior end with caudal process
slightly upturned although damaged in the specimen available. Dorsal margin
384 REVISION OF SOME
with convex central section and steeply sloping antero-dorsal and postero-dorsal
slopes. Greatest length of carapace through the mid-point; greatest height and
width slightly anterior to the mid-point. Left valve larger than the right which it
overlaps along the entire dorsal and ventral margins and overreaches slightly at the
anterior margin. Shell surface finely punctate. Internal features not known.
DIMENSIONS. IN.43506, carapace, length 0-85 mm. ; height 0-59 mm. ; width
0-46 mm.
REMARKS. Bairdia juddiana is sufficiently distinct for the single carapace to be
considered a separate species; it differs from B. hilda in being shorter, higher and more
strongly convex in dorsal view. The postero-dorsal slope of B. hilda is more strongly
concave than that of B. juddiana which lacks the slight anterior upturning of the
antero-dorsal slope seen in the former.
Bairdia jurassica Jones
(PI. 2, figS. I, 2)
1884 Bairdia jurassica Jones : 771, pi. 34, fig. 21.
1884 Bairdia jurassica var tenuis Jones : 771, pi. 34, fig. 22.
DIAGNOSIS. Carapace elongate, drawn out, with acuminate slightly upturned
caudal process. Dorsal margin broadly arched with obliquely sloping antero-dorsal
and postero-dorsal slopes. Left valve larger than the right which it overlaps antero-
dorsally, postero-dorsally and mid-ventrally. Postero-ventrally, the left valve
turned outwards to reveal margin of right valve. Shell surface very finely punctate.
LECTOTYPE. IN.43494, figured Jones 1884, pi. 34, fig. 21, Richmond boring,
depth 1,205 ft.
PARALECTOTYPES. ^.43495, figured Jones 1884, pi. 34, fig. 22, and 10.3617,
carapaces from depth 1,205 ft., Richmond boring.
DESCRIPTION. Carapace drawn out, elongate, greatest length just below mid-
point. Greatest height and width just anterior to mid-point. Dorsal margin
broadly arched with the antero- and postero-dorsal slopes obliquely inclined and
only slightly convex. Anteiior end rounded, posterior end acuminate, not promi-
nently upturned. Ventral margin broadly convex, almost straight. Left valve
larger than the right which it overlaps along the antero-dorsal and postero-dorsal
slopes, especially towards the extreme anterior, and posteriorly along the caudal
process. Left valve overlap prominent mid-ventrally; left valve very noticeably
turned outwards postero-ventrally from the right to reveal the margin of the right
valve. Shell surface finely punctate. Internal features not seen.
DIMENSIONS. IN.43494, carapace, length 0-85 mm. ; height 0-39 mm. ; width
0-31 mm. IN. 43495, carapace, length 0-93 mm. ; height 0-40 mm. ; width 0-34 mm.
10.3617, carapace, length 0-88 mm. ; height 0-43 mm. ; width 0-30 mm.
ENGLISH BATHONIAN OSTRACODA 385
REMARKS. From the dimensions given above it can be seen that the variety
tenuis is more elongate than the other two specimens. Some variants of Bairdia
hilda Jones are similar to the present species which may be distinguished on the
shape of its carapace and the type of overlap by the left valve.
Superfamily CYPRIDACEA Baird 1845
Family PARACYPRIDIDAE Sars 1923
Genus PONTOCYPRELLA Ljubimova 1955
Pontocyprella harrisiana (Jones 1849)
(PL 2, fig. 3)
1849 Bairdia harrisiana Jones : 25, pi. 6, figs. ija-f.
1884 Macrocypris bradiana Jones : 766, pi. 34, fig. 23.
For a full synonymy of Cretaceous forms see Kaye 1965 : 73.
MATERIAL. IN.435oo, left valve figured Jones (1884), depth 1,145 ft. 9 in.-
1,146 ft. 6 in.
REMARKS. Jones (1849 : 25) first recorded this species as Bairdia harrisiana
from a number of horizons, but typically from the Cretaceous (Gault, Leacon Hill;
Chalk, Gravesend and Charlton; Detritus, Charing and from the Speeton Clay,
Yorkshire) .
The presence of this species at a depth of 1,145 ft. 9 in.-i,i46 ft. 6 in. in the
Richmond bore indicates that some contamination of the cored material has occurred.
As Jones himself originally described P. harrisiana, it is somewhat surprising that
he failed to recognize it here.
DIMENSIONS. IN. 43500, left valve, length 0-88 mm.; height 0-42 mm.
Superfamily CYTHERACEA Baird 1850
Family SCHULERIDEIDAE Mandelstam 1959
Subfamily SCHULERIDEINAE Mandelstam 1959
Genus SCHULERIDEA Swartz & Swain 1946
Schuleridea (Schuleridea) jonesiana (Bosquet)
(P.I 2, fig. 5)
1884 Cytheridea subperforata Jones : 772, pi. 34, fig. 26, [not p. 768, pi. 34, fig. 25].
For the synonymy of the Cretaceous forms see Kaye 1964 : 45.
MATERIAL. 1X43490, carapace, figured Jones (1884), Richmond boring, 1,205 ft-
REMARKS. The specimen described by Jones (1884 : 768) fiom 1,151 ft.-
1,151 ft. 6 in. in the Richmond boring is not conspecific with that described from
1,205 ft- The latter is a Cretaceous contaminant.
DIMENSIONS. 1X43490, carapace, length 0-95 mm.; height 0-60 mm.; width
0-43 mm.
386 REVISION OF SOME
Schuleridea (Eoschulerided) subperforata (Jones)
(PI. 2, fig. 4)
1884 Cytheridea subperforata Jones : 768, pi. 34, fig. 25, [not p. 772, pi. 34, fig. 26],
LECTOTYPE. IN.43504, right valve, figured Jones 1884, pi. 34, fig. 25. Richmond
boring, 1,151 ft.- 1,151 ft. 6 in.
REMARKS. Male right valve elongate-oval in outline with a characteristic promi-
nence in the region of the anterior cardinal angle. Anteio-dorsal slope short,
steeply inclined, and concave. Anterior and posterior ends rounded; ventral margin
convex with a prominent antero- ventral incurvature. At least twenty-one anterior
radial pore canals in typical fan-like arrangement. Hinge broken, terminal teeth
incomplete. Duplicature well developed, particularly postero-ventrally. Poor
preservation prevents further description.
In outline this species is similar to the male dimorph of Schuleridea (Eoschuleridea)
bathonica Bate (1967 : 41) but may be distinguished by the antero-dorsal prominence
in the region of the cardinal angle and being slightly more narrowly rounded
posteriorly.
DIMENSIONS. IN.435O4, right valve, length 073 mm. ; height 0-39 mm.
Schuleridea (Eoschuleridea) trigonalis (Jones)
(PI. 2, fig. 7)
1884 Bairdia trigonalis Jones : 767, pi. 34, fig. 19.
DIAGNOSIS. Carapace trigonal in outline with high dorsal margin: Dorsum short,
slightly convex, posteriorly sloping in right valve ; umbonate left valve which promi-
nently overreaches the right. Greatest length of carapace below mid-point.
Greatest height anterior of mid-point in right valve, through mid-point in left.
Anterior end rounded, posterior strongly acuminate. Antero- and postero-dorsal
slopes steeply inclined. Anterior radial pore canals few in number.
HOLOTYPE. IN. 42373, carapace figured Jones 1884, pi. 34, fig. 19; Richmond
boring, depth 1,151 ft.-i,i5i ft. 6 in.
DESCRIPTION. Carapace strongly trigonal in lateral view with the umbonate left
valve projecting noticeably above the right. Posterior end strongly acuminate,
anterior end rounded. Greatest length of carapace below mid-point ; greatest height
of the left valve through mid-point and of the right valve slightly forward of this.
Greatest width median. Ventral margin broadly convex; dorsal margin in the right
valve short and slightly convex, sloping to the posterior. Dorsal margin in the left
valve strongly umbonate. Antero-dorsal and postero-dorsal slopes steeply inclined,
slightly convex. Shell surface very finely punctate. Left valve larger than the right
which it overlaps along the ventral margin, around the posterior and along the
ENGLISH BATHONIAN OSTRACODA 387
antero-dorsal and postero-dorsal slopes. Mid-dorsally the left valve projects strongly
above the right. Internal details not seen. Anterior radial pore canals not clearly
seen but appear to be relatively few.
DIMENSIONS. ^.42373, carapace, length 0-70 mm. ; height 0-48 mm. ; width
0-36 mm.
REMARKS. If the species is dimorphic, the holotype must be that of a female
carapace. A species close to this is Schuleridea (Eoschuleridea] bathonica Bate (1967 :
41) from which it may be distinguished by the strongly umbonate left valve and the
more strongly acuminate posterior.
Genus PRAESCHULERIDEA Bate 1963
Praeschuleridea schwageriana (Jones)
(PI. 2, fig. 6)
1884 Cythere schwageriana Jones : 766, pi. 34, fig. 27.
DIAGNOSIS. Carapace oval in outline with the left valve projecting above the
right. Postero-dorsal slope of left valve rounded, posterior rounded. Shell surface
finely punctate.
HOLOTYPE. ^.43499, female carapace, figured Jones 1884, pi. 34, fig. 27. From
depth 1,145 ft- 9 in.-i,i46 ft. 6 in.
DESCRIPTION. Carapace oval in outline with rounded anterior and posterior
margins. Ventral margin broadly, but not strongly convex in left valve, somewhat
flattened in the right valve. Dorsal margin strongly convex in the left valve,
projecting above the right. Anterior cardinal angle rounded, posterior angle more
sharply acute. Postero-dorsal slope steeply inclined, more strongly convex in the
left valve. Antero-dorsal slope convex in both valves. Greatest length through
mid-point. Greatest height median; greatest width slightly behind mid-point.
Shell surface finely punctate. Left valve larger than the right. No internal
features observed.
DIMENSIONS. 1^43499, female carapace, length 0-60 mm. ; height 0-39 mm. ;
width 0-31 mm.
REMARKS. Jones (1884) referred to but a single specimen for this species and
illustrated a left valve. However, in the plate description he referred to fig. 27 as a
right valve. The actual specimen is in fact a complete carapace, and in the absence
of any evidence to the contrary is considered here to be the holotype.
Praeschuleridea schwageriana is similar to P. subtrigona (Jones & Sherborn 1888)
but is larger and less angular posteriorly : it is close to P. subtrigona intermedia Bate
(1965) but may be distinguished by the more uniformly rounded posterior end.
Given more material showing only slight variation in outline, it might not be possible
to distinguish P. schwageriana from one of the subspecies of P. subtrigona.
388 REVISION OF SOME
Praeschuleridea sp.
(PI. 2, fig. 8)
DESCRIPTION. Carapace trigonal in outline with the left valve strongly umbonate
and projecting above the right valve. Strongly convex when viewed dorsally.
Greatest length, height (left valve) and width pass through mid-point. Right valve
somewhat umbonate in the region of the anterior cardinal angle, below which there
is a deep, oblique groove. Left valve larger than the right. Anterior and posterior
margins rounded; ventral margin broadly convex. Internal details not seen.
Anterior radial pore canals in part seen from the exterior and appear typical for the
genus.
DIMENSIONS. 10.3619, female carapace, length 0-59 mm. ; height 0-43 mm. ;
width 0-35 mm.
REMARKS. This somewhat globose, triangular ostracod is similar in general
outline to Schuleridea trigonalis (Jones 1884) , but differs most markedly in that the line
of greatest length passes through mid-point and not below it as in the latter species.
This species of Praeschuleridea is almost certainly new and was found in a slide of
duplicate material obtained from the Richmond boring; depth 1,151 ft. 6 in. It is
considered inadvisable to erect a new species on only a single specimen.
Family PROGONOGYTHERIDAE Sylvester-Bradley 1948
Subfamily PROGONOCYTHERINAE Sylvester-Bradley 1948
Genus GLYPTOCYTHERE Brand & Malz 1962
Glyptocythere guembeliana (Jones)
(PI. 3, figs, i, 2 ; PI. 4, fig. i)
1884 Cythere guembeliana Jones : 772, pi. 34, figs. 32, 33, [not fig. 31].
1888 Cytheridea pulvinar Jones & Sherborn : 266, pi. 3, figs. za-c.
1888 Cytheridea trapezoidalis Terquem, Jones & Sherborn : 270, pi. 4, figs. la, b.
1967 Glyptocythere guembeliana (Jones) Bate : 49, pi. 13, figs. 10-16, pi. 14, figs. 1-8.
DIAGNOSIS. Carapace subquadrate, elongate in male dimorph. Lateral surface
with transverse ridges extending down from dorsal margin, though generally poorly
developed. Marginal borders compressed. Ventro-lateral margin evenly convex
in female, sharply directed upwards posteriorly in male right valve. Hinge weakly
entomodont.
LECTOTYPE. IN.43493, male right valve from Richmond boring, depth 1,205 ft.
Figured Jones (1884, pi. 34, fig. 33).
PARALECTOTYPE. 10.3338, male carapace from same depth.
REMARKS. This species was revised by Bate (1967 : 49). The specimen illus-
trated is the lectotype (IN. 43493) figured by Jones 1884, pi. 34, fig. 33, and recorded
from a depth of 1,205 ft. The specimen for fig. 32 is missing and that of fig. 31 is
not considered conspecific.
ENGLISH BATHONIAN OSTRACODA 389
DIMENSIONS. Lectotype: IN.43493, male right valve, length 0-96 mm. ; height
0-49 mm.
Genus FASTIGATOCYTHERE Wienholz 1967
Fastigatocy there juglandica (Jones)
(PI. 3, figs. 4, 7, 8; PI. 12, fig. 3)
1884 Cythere juglandica Jones : 766, 768, pi. 34, figs. 36, 37.
1888 Cythere juglandica var. major Jones & Sherborn : 225, pi. 4, figs. 2a-b.
1948 Progonocythere juglandica (Jones) Sylvester-Bradley : 193, pi. 12, figs. 5, 6, pi. 13, fig. 8.
1963 Progonocythere juglandica juglandica (Jones); Grekoff : 1731, pi. 3, fig. 55.
1963 Progonocythere? juglandica (Jones) ; Oertli : pis. 28, 29, 30.
1967 Glyptocythere juglandica (Jones) Bate : 51.
DIAGNOSIS. Carapace sub-rectangular tapering strongly to posterior end. Left
valve larger than right, projecting above right valve dorsally, except mid-dorsally
where the umbonate right valve strongly projects. Ornamentation of transverse
ridges and reticulae radiate down from dorsal margin. Ventro-lateral margins
convex; carapace swollen medially. Flattened marginal borders. Hinge entomo-
dont.
LECTOTYPE. 1X41947, right valve. Richmond boring, 1,146 ft. Figured Jones
(1884).
PARALECTOTYPES. 1X41948-9. Two carapaces from 1,151 ft. 6 in.
REMARKS. This species was described by Sylvester-Bradley (1948) who placed it
in his new genus Progonocythere. Brand & Malz (1962) removed from that genus
those species which have a more quadrate outline, which possess a distinct ornamenta-
tion not usually present in species of Progonocythere s.s., and which also have a dorsal
projection of the dorso-median part of the right valve. The last feature was noted
by Sylvester-Bradley (1948 : 194) and used by Bate (1967 : 51) to justify the assign-
ment of this species to Glyptocythere. Subsequently Wienholz (1967 : 25) separated
from Glyptocythere those species which have a more elongate carapace outline, a more
positive antero-dorsal furrow and strongly diverging ribs radiating from the dorsal
margin. For these species Wienholz erected the genus Fastigatocy there, to which she
assigned juglandica. This assignment is accepted here.
DIMENSIONS. Lectotype : 1X41947, right valve, length 0-93 mm. ; height 047 mm.
1X41948, carapace, length 0-87 mm. ; height 0-51 mm. ; width 049 mm. 1X41949,
carapace, length 0-77 mm. ; height 049 mm. ; width 049 mm.
Genus LOPHOCYTHERE Sylvester-Bradley 1948
REMARKS. There is an understandable reluctance on the part of some ostracod
workers to identify a genus or subgenus simply on an ornamental variation. However,
some ostracod lineages fall naturally into morphological groups in which a particular
ornamental trend is evident. I feel that within the Ostracoda there is ornamentation
390 REVISION OF SOME
of a primary kind which reflects a generic or possibly subgeneric status, and a secon-
dary ornamentation of a more specific character. The latter might be superimposed
upon the former or simply modify it.
Lophocythere clearly contains two distinct morphological groups : the first centred
around Lophocythere ostreata (Jones & Sherborn 1888) with an L-shaped ridge ex-
tending around the anterior margin and bending back along the ventro-lateral margin
as the diagnostic character, and the second around L. bradiana (Jones 1884), in
which several ridges extend across the lateral surface of the carapace.
Whatley (personal communication) informs me that he has, in manuscript, sub-
divided the genus Lophocythere accordingly. As this information will shortly be
published by him it is not my intention to deal further with this generic revision.
Lophocythere bradiana (Jones)
(PI. 3, figs. 3, 5, 6; Text-figs. 3, 4)
1884 Cythere bradiana Jones : 772, pi. 34, figs. $8a-b.
1888 Cytheridea craticula Jones & Sherborn : 272, pi. 4, figs, ga-c, loa-c.
1948 Lophocythere bradiana (Jones) Sylvester-Bradley : 196, pi. 14, figs. 7-10, pi. 15, figs. 8-n.
DIAGNOSIS. Carapace subrectangular, dimorphic. Left valve larger than, and
projects dorsally above, the right. Anterior end broadly rounded, posterior end
triangular with greatest length of carapace through mid-point. Cardinal angles
prominent, rounded. Eye swelling situated at anterior cardinal angle. Lateral
surface ornamented by four major carinae: dorsal carina originates at, or a short
distance from, the anterior margin, curves under the eye swelling, to which a short
offshoot runs, then curves back over dorso-median part of valve to die out below
posterior cardinal angle. The second carina originates at the antero-ventral margin,
runs parallel to the dorsal carina for a short distance then bifurcates, the two
branches running parallel almost to the posterior end where they converge and almost
meet. The fourth carina originates at the antero-ventral margin then follows a
course roughly parallel to lowermost of the two median carinae. A short carina
occurs between the ventral carina and the lowest median carina in the anterior
half of the valve. A second short carina, convex dorsally, occurs between the dorsal
carina and the uppermost of the two median carinae in the posterior half of the valve,
in which region a short, straight carina, is sometimes developed between the two
median carinae. Shell surface reticulate between the carinae.
LECTOTYPE. ^.42372, male carapace, figured Jones (1884), Richmond boring,
depth 1,205 ft.
PARALECTOTYPES. 10.3627-9, one male and two female carapaces from depth
1,205 ft.
DESCRIPTION. Carapace subrectangular, more elongate in the male dimorphs.
Dorsal and ventral margins virtually parallel, with rounded anterior, and triangular
posterior margins. Cardinal angles prominent, rounded, especially prominent in the
larger left valve, the dorsal margin of which projects above the right. Ventral
ENGLISH BATHONIAN OSTRACODA 391
surface ornamented with three longitudinal ridges per valve. Left valve uniformly
overlaps the right along the ventral margin, more strongly so antero-ventrally.
Greatest length of carapace through mid-point, greatest height in the anterior third,
greatest width in the posterior third. Ornamentation of lateral surface as in the
diagnosis. Eye swelling prominently situated below the anterior cardinal angle.
Details of hinge not known from the type material although it is entomodont in
comparative material.
DIMENSIONS. IN.42372, male carapace, length o-6imm.; height 0-32 mm.;
width 0-29 mm. 10.3627, male carapace, length 0-65 mm. ; height 0-34 mm. ; width
0-30 mm. 10.3628, female carapace, length 0-57 mm. height 0-34 mm. ; width
0-29 mm. 10.3629, female carapace, length 0-66 mm. ; height 0-39 mm. ; width
0-34 mm.
REMARKS. Sylvester-Bradley (1948) placed Cytheridea bradiana Jones & Sherborn
in synonymy with Cythere bradiana Jones. However, the specimen described by
Jones & Sherborn is not conspecific but is a paralectotype of Lophocythere septicostata
Bate (1967 : 52, pi. 15, figs. 7-13, pi. 16, figs. 1-4).
FIG. 3. Lophocythere bradiana (Jones). Right valve, complete carapace of lectotype of
Cytheridea bradiana Jones. 11^.42372. x 140.
FIG. 4. Lophocythere bradiana (Jones). Right valve, complete carapace of lectotype of
Cytheridea craticula Jones & Sherborn. 1.1854. x T4°-
392 REVISION OF SOME
Sylvester-Bradley (1948 : 197) considered Cytheridea craticula Jones & Sherborn
1888 to be a subspecies of bradiana because " there is a short anterior stem from
which spring the dorsal, the two median, and the ventral carinae ". Examination
of the types of bradiana and of craticula show this is not, in fact so. The dorsal
carina does not in all specimens reach the anterior margin. Where the reticulate
ornamentation is particularly well developed a cross reticulation gives the impression
that the dorsal carina is joined to the median carina. The ventral carina originates
at the antero-ventral margin, and it is the short carina situated between this and the
lowermost median carina which sometimes joins the stem of the two median carinae.
Although the lectotype of L. bradiana shows the development of a low ridge or
carina between the two median carinae in the posterior part of the valve, many speci-
mens do not show this feature. It is, however, well developed in the male paralecto-
type, 10.3627, and should not be regarded as being a feature of craticula even though
the lectotype of the latter shows it remarkably well.
To summarize, the variations found within a population of Lophocythere bradiana,
such as the occasional imperfect development of the dorsal carina close to the anterior
margin and the presence or absence of a short carina posteriorly between the two
median carinae, do not indicate either a separate species or subspecies, but are part
of the normal variation present in the species.
L. bradiana is followed in the Oxfordian of France by L. multicostata Oertli (1957),
a species which has a similar ornamentation. However, L. multicostata does not
possess the anterior carina situated between the ventral and lower median carina of
bradiana, but has an additional anterior ridge in front of the eye swelling.
The variations within Lophocythere bradiana are illustrated in Text-figs. 3, 4.
Genus MICROPNEUMATOCYTHERE Bate 1963
Micropneumatocythere subconcentrica (Jones)
(PL 4, ng. 7)
1884 Cy 'there subconcentrica Jones : 768, pi. 34, figs. 28, 29.
1967 Micropneumatocythere subconcentrica (Jones) Bate : 60, pi. 21, figs. 1-13.
DIAGNOSIS. Micropneumatocythere with oval carapace tapering posteriorly.
Ventro-lateral margin swollen. Shell surface punctate laterally. Ventral surface
with longitudinal ridges extending on to ventro-lateral margin and turning upwards
anteriorly and posteriorly. Internal details as for genus.
LECTOTYPE. IN.43505, left valve from Richmond boring, depth 1,151 ft. 6 in.
Figured Jones (1884, pi. 34, fig. 28).
REMARKS. This species has been revised by Bate (1967 : 60) and need not be
dealt with fully here. The lectotype, a left valve, is considered to be the specimen
figured by Jones although the caption to the figure states that it is a right valve.
DIMENSIONS. IN.43505, left valve, length 0-56 mm.; height 0-38 mm.
ENGLISH BATHONIAN OSTRACODA 393
Genus TERQUEMULA Blaszyk & Malz 1965
Terquemula blakeana (Jones)
(PI. 4, fig. 8 ; PI. 5, fig. i)
1884 Cytheve blakeana Jones : 772, pi. 34, figs. 34, 35.
1888 Cytheridea transversiplicata Jones & Sherborn : 267, pi. 3, fig. 4.
?i888 Cytheridea blakeana (Jones), Jones & Sherborn : 265, pi. 2, fig. n.
1888 Cytheridea egregia Jones & Sherborn : 267, pi. 3, fig. 5.
1948 Progonocythere blakeana (Jones), Sylvester-Bradley : 191, pi. 12, figs. 3, 4, pi. 13, figs. 4, 5.
1959 Progonocythere ? blakeana (Jones), Triebel & Klingler : 339, 341.
1963 Progonocythere ? blakeana (Jones), Oertli : 36, pi. 25, fig. c, pi. 26, fig. c, pi. 27, fig. c,
pi. 28, fig. c, pi. 30, fig. c.
1965 Terquemula blakeana (Jones), Blaszyk & Malz : 445.
DIAGNOSIS. Carapace oval-elongate, noticeably constricted mid-dorsally and
mid-ventrally. Anterior end rounded, posterior end narrowly rounded. Postero-
lateral part of carapace swollen, overreaching ventral margin. Greatest length
through or slightly above mid-point. Shell surface with large, sunken normal pore
canal openings and a broad reticulation of wrinkles. Muscle scars of Type A (Bate
1963 : 180). Species dimorphic. Hinge entomodont.
LECTOTYPE. 11^.41950, female left valve, Richmond boring, depth 1,205 ft.
Figured Jones (1884).
REMARKS. This species was described fully by Sylvester-Bradley (1948). The
type specimen of Cytheridea egregia Jones & Sherborn, placed by Sylvester-Bradley
in synonymy with blakeana, has been lost, but the original illustration (pi. 3, fig. 5)
suggests that the synonymy is justified.
The specimen described as C. blakeana by Jones & Sherborn could be a juvenile
form of another specimen as suggested by Sylvester-Bradley (1948 : 193).
DIMENSIONS. Lectotype: female left valve, length 0-69 mm. ; height 0-39 mm.
Family CYTHERURIDAE Miiller 1894
Genus METACYTHEROPTERON Oertli 1957
Metacytheropteron drupaceum (Jones)
(PL 5, figs. 4-6)
1884 Cy there drupacea Jones : 772, pi. 34, fig. 30.
I957« Metacytheropteron sp. 50, Oertli : table i.
1963 Cytheropteron jonesanum Cory ell : 854.
1967 Metacytheropteron drupacea (Jones) ; Bate : 44, pi. 10, figs. 1-9.
DIAGNOSIS. Metacytheropteron with elongate/sub-ovate carapace. Greatest
height at anterior cardinal angle, posterior end acuminate. Dorsal margin broadly
convex, especially in .left valve. Shell surface strongly ornamented with triangular
arrangement of longitudinal and obliquely transverse ridges.
GEOL. 17, 8 24
394 REVISION OF SOME
HOLOTYPE. IN. 43498, female carapace from Richmond boring, depth 1,205 ft.
Figured Jones (1884).
REMARKS. Coryell (1963) renamed this species Cytheropteron jonesanum on the
assumption that C. drupacea mentioned by Richter (1867 : 228) and Cy there drupacea
of Jones were homonyms. Reference to Richter (1855 : 529), however, indicates
that the species referred to in 1867 had been placed in Cythereis and not Cy there as
supposed by Coryell. There is, therefore, no nomenclatural problem.
DIMENSIONS. Holotype. 1X43498, carapace, length 0-50 mm. ; height 0-30 mm. ;
width 0-29 mm.
Family TRACHYLEBERIDIDAE Sylvester-Bradley 1948
Genus CYTHEREIS Jones 1849
Cythereis cf. blandu Kaye 1963
(PI. 5, figs. 3, 7)
1884 Cythere (Cythereis) quadrilatera (Roemer), Jones : 766, 772, pi. 34, figs. 39, 40, 41.
REMARKS. Jones recorded Cythereis quadrilatera Roemer from two horizons in
the Richmond boring (1,145 ft. 9 in.-i,i46 ft. 6 in., and 1,205 ft.). These specimens
are probably conspecific with Cythereis blanda Kaye (1963), although the median
and ventro-lateral ridges in Jones's material tend to be more straight and less
strongly developed than in C. blanda. But this may be within the species' variation.
Jones referred to three specimens, here regarded as Cretaceous contaminants, of
Cythere (Cythereis) quadrilatera but only two now remain ; these are a complete cara-
pace, IN.4349I, figured Jones 1884, pi. 34, fig. 40, and a single left valve IN.43502,
figured pi. 34, fig. 39. The specimen illustrated by fig. 41 is missing and the illustra-
tion suggests that this latter specimen was not conspecific with the other two.
DIMENSIONS. IN.4349I, carapace, length 0-77 mm. ; height 0-46 mm. ; width
0-32 mm. IN.435O2, left valve, length 0-66 mm. ; height 0-43 mm.
Family and Genus uncertain
" Cythere " tenella Jones
(PI. 5, fig- 8)
1884 Cythere tenella Jones : 772, pi. 34, fig. 24.
REMARKS. This small, poorly-preserved ostracod is probably a juvenile instar.
HOLOTYPE. 1X43492, complete carapace, Richmond boring, depth 1,205 ft-
DIMENSIONS. Length 0-38 mm. ; height 0-21 mm. ; width 0-15 mm.
III. THE T. R. JONES & C. D. SHERBORN COLLECTION
AND THE H. H. WINWOOD COLLECTION
The following figured specimens of species described by Jones & Sherborn (1888)
are no longer represented in their collection and may be presumed lost. Only the
first five are known to have been presented to the Museum.
ENGLISH BATHONIAN OSTRACODA 395
Bairdia juddiana Jones, p. 253.
Cytheridea bicarinata Jones & Sherborn, p. 270, pi. 4, figs. 5#-c.
Cytheridea renoides Jones & Sherborn, p. 266, pi. 3, figs. la-c.
Cytheridea winwoodiana Jones & Sherborn, p. 259, pi. I, figs. 2a-d.
Cytheridea eximia Jones & Sherborn, p. 273, pi. 5, figs. 5«-c.
Bairdia trigonalis Jones, p. 253.
Bythocypris winwoodiana Jones & Sherborn, p. 252, pi. 5, figs. xa-c.
Cythere juglandica var. minor Jones & Sherborn, p. 255, pi. 4, figs. 3#-c.
Cythere ? speciosa Jones & Sherborn, p. 254, pi. 2, figs. loa-c. (This species
was recorded as lost by Jones & Sherborn, p. 254.)
Cythere trapezioides Jones & Sherborn, p. 256, pi. 5, figs. ioa-c.
Cythere walfordiana Jones & Sherborn, p. 255, pi. 5, figs. ga-c.
Cytheridea egregia Jones & Sherborn, p. 267, pi. 3, figs. 5«-c.
Cytheridea politula Jones & Sherborn, p. 265, pi. 5, figs. ja-c.
Cytheridea pura Jones & Sherborn, p. 269, pi. 3, figs. na-d.
Cytheridea retorrida Jones & Sherborn, p. 260, pi. i, figs. 8a~c.
Cytheridea rugifera Jones & Sherborn, p. 271, pi. 5, figs. na-c.
Cytheridea sedata Jones & Sherborn, p. 261, p. i, figs. loa-c.
Cytheridea subeminula Jones & Sherborn, p. 261, pi. 5, figs. 8a-c.
Cytheridea ventrosa Jones & Sherborn, p. 269, pi. 3, figs. loa-c.
Cythereis walfordiana Jones & Sherborn, p. 257, pi. 5, figs. I2a-c.
Order PODOCOPIDA Miiller 1894
Suborder PLATYCOPINA Sars 1866
Family GYTHERELLIDAE Sars 1866
Genus CYTHERELLA Jones 1849
Cytherella fullonica Jones & Sherborn
(PI. 5, ng. 9; PI. 6, fig. i)
1888 Cytherella fullonica Jones & Sherborn : 274, pi. i, figs. I2a—c.
1963 Cytherella fullonica Jones & Sherborn; Bate : 184, pi. i, figs, i, 2.
DIAGNOSIS. Carapace subrectangular with characteristically steep postero-dorsal
slope. Shell surface smooth, with dorso-median muscle-scar depression.
LECTOTYPE. 1.1857, left valve, figured Jones & Sherborn 1888, from the Blue
Fuller's Earth Clay.
PARALECTOTYPES. 1.4004 (Jones & Sherborn Collection); 10.3636-41 (Winwood
Collection), left and right valves. Three specimens from the Blue, and four from
the Yellow Fuller's Earth Clay.
REMARKS. This species has been previously described in Bate (1963 : 184).
DIMENSIONS. Lectotype: 1.1857, kft valve, length 0-64 mm.; height 0-34 mm.
Paralectotype lo. 4004, left valve, length 0-58 mm.; height 0-32 mm.
396 REVISION OF SOME
Genus CYTHERELLOIDEA Alexander 1929
Cytherelloidea catenulata (Jones & Sherborn)
(PI. 6, figs. 2, 3)
1888 Cytherella catenulata Jones & Sherborn : 274, pi. 5, figs. 6a-c.
1948 ? Cytherelloidea catenulata (Jones & Sherborn) Sylvester-Bradley : 200, pi. 14, fig. n.
1963 Cythetelloidea catenulata (Jones & Sherborn); Bate : 184, pi. i, figs. 3-6.
1963 Cytherelloidea catenulata (Jones & Sherborn) ; Oertli : 37, pis. 27, 29.
1964 Cytherelloidea catenulata (Jones & Sherborn) ; Bate : 8.
DIAGNOSIS. Carapace with crescent-shaped swelling situated close to posterior
margin of valve. Dorsal limb of crescent bends over and curves forwards close to
ventral margin to produce a swelling shaped like a question mark. Shell surface
ornamented with longitudinal and terminal (parallel to margins) striae producing a
reticulation.
LECTOTYPE. 1.1876, left valve, figured Jones & Sherborn, from the Blue Fuller's
Earth Clay.
PARALECTOTYPES. 1.1846 (left valve) and 10.3525 (right valve), Jones & Sherborn
Collection; 10.3649-53, left and right valves from the Winwood Collection. All
specimens from the Blue Fuller's Earth Clay.
REMARKS. This species was described by Bate (1963 : 184-5).
DIMENSIONS. 1.1876, left valve, length 0-68 mm.; height 0-37 mm. 1.1846,
left valve; length 0-68 mm. ; height 0-36 mm. 10.3525, right valve, length 0-68 mm. ;
height 0-38 mm.
Cytherelloidea refecta (Jones & Sherborn)
(PI. 6, figs. 4-7)
1888 Cytheridea refecta Jones & Sherborn : 262, pi. 2, figs. 3«, b.
DIAGNOSIS. Carapace oval, more elongate in male dimorph. Anterior and
posterior margins rounded. Posterior cardinal angle prominent, postero-dorsal slope
slopes steeply to posterior. Central part of valve with broad depression bounded
below by broad crescentic swelling. Crescentic furrow delimits lower surface of
swelling. A small rounded swelling is situated to the inside of the anterior part of
crescentic swelling.
LECTOTYPE. 1.1850, female right valve, figured Jones & Sherborn (1888), from
the Blue Fuller's Earth Clay.
PARALECTOTYPES. 10.3931-2, female left valve and male left valve from the
Winwood Collection. Both specimens from the Blue Fuller's Earth Clay.
DESCRIPTION. Carapace oval in outline with rounded anterior and posterior
margins and prominent posterior cardinal angle. The postero-dorsal slope is steeply
inclined, but slightly convex. Dorsal margin of both valves slightly concave antero-
ENGLISH BATHONIAN OSTRACODA 397
medially. Ventral margin convex in the right valve, concave in the left. Shell
surface with a broad, deep furrow, crescentic in shape which commences postero-
dorsally and terminates antero-dorsally. Above this furrow a broad swelling is
developed, also crescentic but rather irregular in outline. Anteriorly the swelling
appears to turn back sharply upon itself for a short distance before dying out. To
the inside of this swelling and situated in the angle of the anterior turn-back, a small
circular swelling is developed. The main crescentic swelling is delimited on its
dorsal side by a broad central depression. In the region of the anterior margin the
surface of the shell may be seen to be weakly reticulate. Internally the dorsal edge
of the right valve possesses a groove for the reception of the valve margin of the
smaller left valve.
DIMENSIONS. 1.1850, female right valve, length 0-60 mm.; height 0-26 mm.
10.3931, female left valve, length 0-52 mm. ; height 0-28 mm. 10.3932, male left
valve, length 0-71 mm. ; height 0-38 mm.
REMARKS. Cytherelloidea refecta has a distinctive ornamentation which separates
it from other Jurassic species of the genus. Within the Jurassic the predominant
ornamentation of the cytherelloideas consists of either a peripheral ridge or swelling
and/or a central ridge or swelling. The development of strong lateral ridges extending
from the posterior region, though present in some Jurassic species (Field 1966), is
predominantly a feature of Cretaceous and Tertiary species.
Suborder PODOCOPINA Sars 1866
Superfamily BAIRDIACEA Sars 1888
Family BAIRDIIDAE Sars 1888
Genus BAIRDIA McCoy 1844
Bairdia Hilda Jones
(PI. 4, fig- 5)
1884 Bairdia hilda Jones : 771, pi. 34, fig. 20.
1888 Bairdia fullonica Jones & Sherborn : 253, pi. 5, figs. ^a-c.
For complete synonymy see p. 383.
REMARKS. Sylvester-Bradley (1948 : 199) first pointed out that the variation
found within specimens of Bairdia hilda and Bairdia fullonica was such that there
was as yet no evidence for regarding these species as distinct. Accordingly they
were placed in synonymy. The lectotype of B. fullonica (1.1873, a right valve from
the Blue Fuller's Earth Clay, figured Jones & Sherborn) was in the Jones & Sherborn
Collection, whilst the paralectotype, also a right valve (10.3554) and from the Blue
Fuller's Earth Clay, was in the Win wood Collection.
DIMENSIONS. 1.1873, right valve, length 079 mm.; height 0-42 mm. 10.3554,
right valve, length 0-76 mm. ; height 0-39 mm.
398 REVISION OF SOME
In addition to the types there are four right valves and one left valve (1.1848 from
the Jones & Sherborn Collection and 10.3911-2 and 10.271 from the Winwood
Collection) from the Blue Fuller's Earth Clay and referred to by Jones & Sherborn
on p. 253.
Bairdia sherborni sp. nov.
(PI. 6, fig. 8; PL 7, fig. i)
DIAGNOSIS. Carapace sub-rectangular in outline. Dorsal margin very slightly
convex, sloping slightly to posterior. Ventral margin strongly incurved medially.
Anterior end obliquely rounded. Posterior end narrowly rounded, somewhat
tapering, but not upturned. Left valve larger than the right. Muscle scars as for
the genus. Shell surface strongly and coarsely pitted.
HOLOTYPE. 10.3913, complete carapace from the Yellow Fuller's Earth Clay,
Winwood Collection.
PARATYPE. 10.3557, complete carapace from the Fuller's Earth Clay, Mockler
Collection.
DESCRIPTION. Carapace sub-rectangular, elongate bairdioid in outline, the dorsal
margin very slightly convex but almost straight, sloping gently to the posterior.
The ventral margin is typical of the genus, being strongly incurved medially.
Anteriorly the margin is obliquely rounded, sharply cut just above a line running
through mid-point (line of greatest length) by the inclined antero-dorsal slope.
Postero-dorsal slope slightly concave, postero-ventral slope convex. Posterior end
tends to be tapered without the strong upturning of most bairdias. Greatest height
of carapace through the anterior cardinal angle ; greatest width in the posterior third.
Shell surface distinctly and evenly pitted. Left valve larger than the right which it
overlaps mid-ventrally though not antero-ventrally and postero-ventrally. Antero-
dorsally and postero-dorsally the gape is replaced by a prominent overlap of the
right valve by the left. Internal details not observed apart from the muscle scars
which may be observed as impressions on the internal cast of the holotype. They
comprise seven adductor scars arranged in a circle with an eighth situated at the
centre and a ninth, probably also an adductor scar, situated dorsally to this group.
An additional dorsal scar is placed above that last mentioned whilst antero-ventrally
there are two oval " mandibular " scars and antero-dorsally a single " antennal "
scar.
DIMENSIONS. 10.3913, carapace, length 0-58 mm. ; height 0-31 mm. ; width
0-21 mm. 10.3557, carapace, length 0-71 mm.; height 0-39 mm.; width 0-27 mm.
REMARKS. Only two specimens have been found, one in each of two major
collections. The outline of the carapace; obvious antero- ventral and postero-ventral
gape and distinct surface pitting distinguish this species easily from all other Jurassic
bairdiids.
ENGLISH BATHONIAN OSTRACODA 399
Superfamily GYPRIDACEA Baird 1845
Family PARACYPRIDIDAE Sars 1923
Genus PARACYPRIS Sars 1866
Paracypris terraefullonicae (Jones & Sherborn)
(PI. 7, figs. 2, 4)
1888 Macrocypris terrae-fullonicae Jones & Sherborn : 252, pi. 5, figs. $a-c.
1888 Macrocypris horatiana Jones & Sherborn : 252, pi. 5, figs. za-c.
1967 Paracypris terraefullonica (Jones & Sherborn) Bate : 27, pi. i, figs. 1-6.
DIAGNOSIS. Carapace elongate, posteriorly acuminate. Anterior end rounded.
Ventral margin almost straight in the larger left valve, more strongly concave in the
right. Dorsal margin arched with antero-dorsal slope tending to be slightly concave,
more noticeably so in the right valve. Shell surface smooth. Anterior and posterior
vestibules well-developed. Radial pore canals branching.
LECTOTYPE. 1.1875, left valve, figured Jones & Sherborn 1888, pi. 5, figs. 3«-c,
from the Blue Fuller's Earth Clay.
OTHER MATERIAL. 1.1874, right valve (lectotype of Macrocypris horatiana Jones
& Sherborn), figured Jones & Sherborn 1888, pi. 5, figs. 2a-c, and 10.3548-53 (para-
lectotypes of M. horatiana and M. terraefullonicae], from the Blue Fuller's Earth
Clay.
REMARKS. Paracypris terraefullonicae has been fully described in Bate (1967 : 27).
The additional specimens listed above (10.3548-53) were found in the Winwood
Collection and are paralectotypes of Macrocypris horatiana and Macrocypris terrae-
fullonicae which " species " are the right and left valves respectively of Paracypris
terraefullonicae.
DIMENSIONS. 1.1875, left valve, length 0-60 mm.; height 0-28 mm. 1.1874,
right valve, length 0-58 mm. ; height 0-26 mm.
Superfamily CYTHERACEA Baird 1850
Family BYTHOCYTHERIDAE Sars 1926
Genus MONOCERATINA Roth 1928
Monoceratina visceralis (Jones & Sherborn)
(PI- 7> fig- 3)
1888 Cytheridea visceralis Jones & Sherborn : 268, pi. 3, figs. 6a-c.
DIAGNOSIS. Carapace sub-rectangular in outline with rounded anterior end; the
antero-dorsal margin tends to curve obliquely backwards. Posterior end triangular
with greatest prolongation situated in dorsal half of valve. Dorsal and ventral
margins parallel. A distinct median sulcus present in dorsal half of valve only.
400 REVISION OF SOME
Valve convex, prominently swollen in postero-ventral region. Shell surface strongly
pitted. Postero-ventral margin slightly serrated. Hinge with strong, straight,
median bar.
HOLOTYPE. 1.1830, left valve, figured Jones & Sherborn 1888, from the Blue
Fuller's Earth Clay.
DESCRIPTION. The carapace is sub-rectangular in outline with a triangular
posterior end, the ventral margin of which is longer than the straight postero-dorsal
margin, and is evenly serrated along its length. At the postero-ventral angle there
is a distinct convexity of the margin. Anterior cardinal angle prominently rounded.
Dorsal and ventral margins parallel. A distinct vertical median sulcus is present in
the dorsal half of the valve only. Carapace strongly swollen along the ventro-lateral
border and postero-ventrally where the swelling is strongly undercut. Shell surface
coarsely pitted, the pits extending uniformly over the valve. Line of greatest length
lies above mid-point, whilst the line of greatest height passes approximately through
the centre of the valve. Internally the hinge has been damaged, only the posterior
part of the strong hinge bar being present. No other internal details observed.
DIMENSIONS. 1.1830, left valve, length 0-66 mm.; height 0-30 mm.
REMARKS. Only the holotype is known, but it is distinct from the other species
of Monoceratina described from the Jurassic. M. visceralis is close to M. vulsa
(Jones & Sherborn), although the latter has a much coarser ornamentation, almost
reticulate, whilst the median sulcus is much more strongly developed, as is the
under-cutting of the ventro-lateral and postero-ventral swelling. M. vulsa is also
distinguished by possessing a distinct furrow paralleling the anterior margin.
M. ungulina Triebel & Bartenstein (1938, pi. I, figs. 3, 4) may be distinguished on
outline, the posterior extremity being situated higher up on the valve because of the
shorter postero-dorsal slope. The anterior margin is also uniformly rounded with
a flattened marginal border.
Monoceratina vulsa (Jones & Sherborn)
(H. 7, ng. 5)
1888 Cytheridea vulsa Jones & Sherborn : 263, pi. 2, figs. $a, b.
1938 Monoceratina vulsa (Jones & Sherborn) Triebel & Bartenstein : 516, pi. 3, figs. 17, 18.
1960 Monoceratina cf. vulsa (Jones & Sherborn) ; Lutze : 433, pi. 37, figs. $a, b.
?I963 Monoceratina sp. juv. aff. vulsa (Jones & Sherborn); Plumhoff : 48, pi. n, figs. 167, 168.
1963 Monoceratina vulsa (Jones & Sherborn); Bate : 189, pi. 3, figs. 5-12.
19630 Monoceratina vulsa (Jones & Sherborn); Bate : 26, pi. i, fig. 6.
1964 Monoceratina vulsa (Jones & Sherborn) ; Bate : 9.
DIAGNOSIS. Shell sub-rectangular in lateral view, convex in dorsal view. Vertical
median sulcus deeply incised, surrounded below and to the sides by a prominent
swelling which is strongly undercut ventro-laterally. Lateral swelling separated
from anterior margin by a furrow which runs parallel to that margin. Shell surface
strongly pitted, the raised borders of the pits giving the surface a wrinkled appearance.
ENGLISH BATHONIAN OSTRACODA 401
LECTOTYPE. 1.1842, a badly damaged left valve, figured Jones & Sherborn 1888,
from the Blue Fuller's Earth Clay.
REMARKS. Only a single damaged valve remains of the two specimens originally
placed in this species by Jones & Sherborn (p. 263), and this is the figured specimen.
A complete description was given by Bate (1963) who incorrectly recorded the
lectotype as coming from the Richmond boring.
DIMENSIONS. 1.1842, length 0-64 mm.; height 0-32 mm. Both measurements
would have been slightly larger if the specimen had not suffered slight damage
posteriorly and along the dorsal margin.
Family TRACHYLEBERIDIDAE Sylvester-Bradley 1948
Subfamily TRACHYLEBERIDINAE Sylvester-Bradley 1948
Genus OLIGOCYTHEREIS Sylvester-Bradley 1948
Oligocythereis fullonica (Jones & Sherborn)
(PI. 7, fig. 6)
1888 Cythereis fullonica Jones & Sherborn : 256, pi. 4, figs,
1948 Cythereis cf. fullonica Jones & Sherborn; Sylvester-Bradley : 186, pi. 12, figs. 7, 8 [not
figs. 9, 10], pi. 13, fig. 3 [not fig. 9].
I948« Oligocythereis fullonica (Jones & Sherborn) Sylvester-Bradley : 796.
1963 Oligocythereis fullonica (Jones & Sherborn) ; Oertli : 39, pi. 25, fig. a?, pi. 29, fig. a, [not
pi. 24, fig. a, pi. 26, fig. a].
1967 Oligocythereis fullonica (Jones & Sherborn); Bate : 61, pi. 21, fig. 14, [not fig. 15].
DIAGNOSIS. Carapace subquadrate in outline with a broadly rounded anterior and
broadly triangular posterior end. Anterior and posterior margins with small
denticles. Ventral surface flattened, V-shaped in outline; ventral margin gently
convex anteriorly and posteriorly. Dorsal margin almost straight, approximately
paralleling ventral margin. Anterior cardinal angle extremely prominent with a
large, rounded eye tubercle below. A short, thick ridge extends obliquely antero-
ventrally from eye tubercle. Dorso-median ridge extends back from eye tubercle,
enlarging posteriorly to give carapace triangular outline in dorsal view. Termination
of dorso-median ridge produces a sharply pointed tubercle, angular in outline, the
two sides of which are at right angles. Postero-ventrally a short, raised ridge almost
vertically directed bends round at an oblique angle to extend forwards ventro-laterally
bending upwards antero-ventrally to terminate in a prominent tubercle. This
ventro-lateral ridge describes a broadly crescentic outline around a prominent,
irregular tubercle. Prominent tubercle developed slightly anterior of mid-point.
Shell surface very finely punctate. Left valve larger than right.
LECTOTYPE. 1.1871, complete carapace, figured Jones & Sherborn 1888, from the
Blue Fuller's Earth Clay.
DESCRIPTION. Carapace subquadrate in outline, ornamented as diagnosed. The
angular outline of the shell is produced dorsally by the prominent antero-dorsal
402 REVISION OF SOME
eye tubercle and the strong postero-dorsal tubercle and ventrally by the postero-
ventral and antero- ventral terminations of the ridge which extends along the ventro-
lateral margin. The large tubercle situated slightly anterior of mid-point is the
muscle scar node characteristic of the family. The anterior and posterior margins
possess small denticles whilst the marginal borders bear a few additional nodes.
Shell surface finely punctate. In dorsal view the outline of the carapace in the
median part is V-shaped as is the ventral surface. This outline is produced by the
dorso-median and ventro-lateral ridges diverging slightly towards the posterior end.
Left valve larger than the right which it overlaps along the ventral margin and in
the region of the cardinal angles and overreaches along the postero-dorsal slope.
Internal details not observed in the type material, but the entomodont type hinge
has been described by Sylvester-Bradley (1948 : 187).
DIMENSIONS. Lectotype, 1.1871, carapace, length 0-54 mm.; height 0-32 mm.;
width 0-27 mm.
REMARKS. Sylvester-Bradley (1948 : 187) noted variation within the ornamenta-
tion of specimens which he placed in 0. fullonica (Jones & Sherborn), and it has been
current practise to retain these within the species. The forms possessing a postero-
dorsal tubercle from which three arms radiate (see Sylvester-Bradley 1948, pi. 12,
figs. 9, 10) should be assigned to a new species. 0. fullonica should be retained only
for those specimens in which the postero-dorsal tubercle has two arms at right angles,
and in which the muscle scar node to the centre of the valve is completely isolated
and not joined to the anterior termination of the ventro-lateral ridge.
Family SCHULERIDEIDAE Mandelstam 1959
Subfamily SCHULERIDEINAE Mandelstam 1959
Genus SCHULERIDEA Swartz & Swain 1946
Subgenus EOSCHULERIDEA Bate 1967
Schuleridea (Eoschuleridea) horatiana (Jones & Sherborn)
(PI. 7, fig. 7; Text-fig. 5)
1888 Cytheridea horatiana Jones & Sherborn : 263, pi. 2, figs. $a, b.
DIAGNOSIS. Carapace oval/elongate, anteriorly rounded, posteriorly tapering,
narrowly rounded. Greatest length of carapace below mid-point, greatest height
at anterior cardinal angle. Elongate eye swelling with groove beneath present in
right valve below anterior cardinal angle. Left valve projects slightly above right
just behind anterior cardinal angle. Shell surface punctate with widely scattered,
large, normal pore canals. Twenty-five to thirty radial pore canals splayed fan-like
around anterior margin. Muscle scars type C. Hinge paleomerodont.
LECTOTYPE. 1.1852, male right valve, figured Jones & Sherborn 1888, from the
Blue Fuller's Earth Clay.
PARALECTOTYPE. 10.3940, male carapace from Blue Fuller's Earth Clay, Winwood
Collection.
ENGLISH BATHONIAN OSTRACODA 403
DESCRIPTION. Carapace oval/elongate in outline with uniformly rounded anterior
margin and narrowly rounded, tapered posterior. The line of greatest length passes
well below the mid-point. Greatest height at the anterior cardinal angle, greatest
width just behind the mid-point. Dorsal margin convex, slightly umbonate in the
left valve just behind the anterior cardinal angle. Ventral margin convex with an
antero-median incurvature. Shell surface punctate with widely scattered, large,
FIG. 5. Muscle scars, x 200. Schuleridea (Eoschuleridea) horatiana (Jones & Sherborn),
lectotype, 1.1852.
rounded, normal pore canals. An elongate eye swelling is situated below the
anterior cardinal angle in the right valve and has an oblique furrow situated beneath.
Muscle scars as viewed externally of type G (Bate 1963). Anterior radial pore
canals splayed fan-like around the anterior margin, largely concentrated below
mid-height, approximately twenty-five to thirty in number. Hinge paleomerodont
as seen in the right valve. Terminal teeth larger anteriorly, where there are five,
than posteriorly, where there are eight. Median bar smooth. Left valve larger
than right. Inner margin and line of concrescence coincide, anterior duplicature
broad.
DIMENSIONS. 1.1852, right valve, length 0-85 mm.; height 0-46 mm. 10.3940,
carapace, length 0-79 mm. ; height 0-46 mm. ; width 0-36 mm.
REMARKS. Both available specimens of this species are male dimorphs as indi-
cated by their shell outline. No female dimorph has been recognized. This species
is very close to the male dimorph of Schuleridea (Eoschuleridea) bathonica Bate (1967)
although the latter has fewer anterior radial pore canals and does not have the line
of greatest length as ventrally positioned as in horatiana. Male dimorphs of
Schuleridea species are notoriously alike and extremely difficult to tell apart. Until
404 REVISIONOFSOME
a female dimorph has been identified for horatiana from the Fuller's Earth it is
preferred to regard these two species as distinct.
Genus ASCIOCYTHERE Swain 1952
Asciocythere obovata (Jones & Sherborn)
(PI. 7, fig. 8; PI. 8, figs. 2, 3, 7; Text-fig. 6)
1888 Cytheridea obovata Jones & Sherborn : 264, pi. 2, figs. 6a-c.
DIAGNOSIS. Carapace oval in lateral view, strongly convex in dorsal view.
Dorsal margin broadly arched, curving down into well-rounded anterior and more
narrowly rounded posterior margin without change in slope at cardinal angles.
Ventral margin broadly convex. Line of greatest length slightly below mid-point.
Shell surface very finely punctate.
LECTOTYPE. 1.1836, right valve, figured Jones & Sherborn 1888, pi. 2, figs. 6a-c,
from the Yellow Fuller's Earth Clay.
PARALECTOTYPE. 10.3938, carapace from the Yellow Fuller's Earth Clay,
Winwood Collection.
OTHER MATERIAL. 10.3939, left valve from the Blue Fuller's Earth Clay, Winwood
Collection.
DESCRIPTION. Carapace oval in outline with strongly arched dorsal margin and
indistinct cardinal angles, the dorsal margin curving down without a break into the
rounded anterior margin and the more narrowly rounded posterior margin. Ventral
margin broadly convex. Shell surface appears smooth but is in fact very finely
punctate. Greatest length of carapace just below mid-point. Greatest height
median; greatest width just behind mid-point. Left valve larger than right, over-
lapping the right along the ventral, posterior and postero-dorsal slopes. Antero-
dorsally and anteriorly the valves tend to gape and there is no overlap. Hinge
antimerodont : right valve with ten dorsally bifid anterior teeth and nine dorsally
bifid posterior teeth. Median groove very short and loculate. Left valve hinge not
seen. Inner margin and line of concrescence coincide, duplicature strongly
developed. Anterior radial pore canals widely spaced and only very slightly
curved, almost straight, sixteen in number. Muscle scars not observed.
DIMENSIONS. 1.1836, right valve, length 0-54 mm.; height 0-34 mm.; 10.3938,
carapace, length 0-49 mm. ; height 0-35 mm. ; width 0-30 mm. 10.3939, left valve,
length 0-62 mm. ; height 0-42 mm.
REMARKS. Only two species of Asciocythere have been described from the British
Middle Jurassic so far; A. lacunosa Bate 1963^ and A. acuminata Bate 1964, both
from the Bajocian. The present species, the first to be described from the Bathonian,
is not so posteriorly acuminate as A. acuminata and not so strongly pitted as
A. lacunosa.
ENGLISH BATHONIAN OSTRACODA 405
FIG. 6. Asciocythere obovata (Jones & Sherborn). Internal view, right valve, lectotype,
1.1836. x 115.
Genus PRAESCHULERIDEA Bate 1963
Praeschuleridea subtrigona subtrigona (Jones & Sherborn)
(PL 8, figs. 4-6)
1888 Cytheridea subtrigona Jones & Sherborn : 265, pi. 2, figs. ga-c.
1963 Praeschuleridea subtrigona (Jones & Sherborn) Bate : 207, pi. 12, figs. 12-16, pi. 13,
figs. 1-9.
1963(3 Praeschuleridea subtrigona (Jones & Sherborn); Bate : 41.
1964 Praeschuleridea subtrigona subtrigona (Jones & Sherborn); Bate : 22.
1965 Praeschuleridea subtrigona subtrigona (Jones & Sherborn); Bate : 124.
DIAGNOSIS. Carapace oval-subtrigonal in outline, punctate. Length of adult
of the order of (female) 0-56 mm. ; (male) 0-58 mm.
LECTOTYPE. 1.1838, male right valve, not the figured specimen of Jones &
Sherborn, which was of a left valve. Blue Fuller's Earth Clay.
PARALECTOTYPES. 10.3935-7, female and two male carapaces, Blue Fuller's
Earth Clay, Win wood Collection.
REMARKS. Praeschuleridea subtrigona is the type species of the genus and as such
has been described fully in earlier publications. Three subspecies are recognized
so far, subtrigona subtrigona being the smallest and having the longest stratigraphical
range: Middle Bajocian (Sonninia sowerbyi Zone, Hyperlioceras discites Subzone) to
Upper Bathonian (Prohecticoceras retrocostatum Zone) . The other two subspecies, P.
subtrigona magna Bate (1964) and P. subtrigona intermedia Bate (1965), have much
shorter ranges and are restricted to the Bajocian. The measurements of the type
specimens have not previously been given and are included here.
DIMENSIONS. 1.1838, male right valve, length 0-53 mm.; height 0-32 mm.
10.3935, male carapace, length 0-55 mm. ; height 0-37 mm. ; width 0-29 mm. 10.3936,
male carapace, length 0-54 mm.; height 0-36 mm.; width 0-29 mm. 10.3937,
female carapace, length 0-50 mm. ; height 0-35 mm. ; width 0-30 mm. Comparison
of the above measurements with those given in Bate (1963 : 209) shows that the size
of the P. subtrigona subtrigona has remained stable since the Bajocian.
406 REVISION OF SOME
Genus EOCYTHERIDEA Bate 1963
Eocytheridea sp.
(PI. 8, fig. i)
REMARKS. Known from a single right valve, possibly of a male dimorph
(Winwood Collection). The surface of the valve is strongly pitted by rather broad
pits which, towards the centre of the valve, produce a reticulation but in the antero-
dorsal and postero-dorsal areas are elongated obliquely towards the centre of the
dorsal margin. This elongation produces a series of ridges which radiate down from
the centre of the dorsal margin. The ornamentation readily distinguishes this species
from Eocytheridea faveolata Bate (1964) which has a more uniform reticulate orna-
mentation without the development of obliquely radiating ridges in the dorsal part
of the carapace. The hinge is hemimerodont. Twelve long, slightly curved anterior
radial pore canals pass through the broad duplicature.
This is the first record of the genus outside the Bajocian
DIMENSIONS. 10.3906, right valve, length 0-77 mm. ; height 0*36 mm.
Family CYTHERIDEIDAE Sars 1925
Subfamily CYTHERIDEINAE Sars 1925
Genus HADROCYTHERIDEA nov.
DERIVATION OF NAME, hadros, Gr., well developed + cytheridea.
DIAGNOSIS. Carapace robust, expanded posteriorly in both width and height.
Hinge straight, antero-dorsal and postero-dorsal slopes steeply angled. Anterior
end rounded, posterior end triangular in elongate male dimorph. Female with
triangular posterior end, slightly upturned. Line of greatest length lower in female
dimorph. Postero-ventral part of carapace characteristically swollen, particularly
in the female. Hine antimerodont. Inner margin and line of concrescence coincide;
duplicature broad. Anterior radial pore canals long, straight, widely and uniformly
spaced, nine in the type species. In front of a vertical row of four small, oval
adductor scars, there is a bill-hook or broadly heart-shaped antero-dorsal antennal
scar and a small antero-ventral mandibular scar. Left valve larger than the right.
Shell surface coarsely pitted.
TYPE SPECIES. Cytheridea dolabra Jones & Sherborn 1888.
REMARKS. This new genus has been placed in the Cytherideidae on the basis of
carapace morphology and muscle scar pattern. It is easily recognizable by its
rectangular/quadrate outline, postero-ventral convexity and position of greatest
height and width towards the posterior. The outline of the male dimorph is to some
extent reminiscent of Fabanella Martin 1961, but that of the female is completely
different.
ENGLISH BATHONIAN OSTRACODA
Hadrocytheridea dolabra (Jones & Sherborn)
(PI. 9, figs. 1-8; PI. 10, fig. i; Text-fig. 7)
407
1888 Cytheridea dolabra Jones & Sherborn : 267, pi. 3, figs.
1888 Cytheridea puteolata Jones & Sherborn. : 259, pi. i, figs. ja-c.
1888 Cytheridea parallela Jones & Sherborn : 260, pi. i, figs. ga-c.
1888 Cytheridea pentagonalis Jones & Sherborn : 261, pi. 2, figs. la-c.
1888 PCytheridea ignobilis Jones & Sherborn : 268, pi. 3, figs. ga-c.
DIAGNOSIS. Hadrocytheridea with robust dimorphic carapace. Shell surface
finely punctate between distinct circular pits, at the centre of which is a large,
circular normal pore canal opening. Other details as for genus.
LECTOTYPE. 1.1851, female right valve, figured Jones & Sherborn 1888, pi. 3,
figs. 3«-c. Blue Fuller's Earth Clay, Midford.
PARALECTOTYPES. 1.1844, male left valve, figured Jones & Sherborn 1888, pi. i,
figs. ja-c. This specimen was described by Jones & Sherborn as Cytheridea puteolata
and is selected here as lectotype of that species. Additional paralectotypes of
C. puteolata are 10.3921-2 from the Winwood Collection.
1.1859, ma-le carapace, figured Jones & Sherborn 1888, pi. i, figs. ga-c. This was
described as Cytheridea parallela and is here selected as lectotype of that species.
Additional paralectotypes of C. parallela are 10.3924-8 from the Winwood Collection.
1. 1866, female left valve, figured Jones & Sherborn 1888, pi. 2, figs. la-c. This is
the holotype of Cytheridea pentagonalis.
All the above paralectotypes of Hadrocytheridea dolabra apart from 1. 1866, which
comes from the Yellow Fuller's Earth Clay, come from the Blue Fuller's Earth Clay
of Midford, near Bath.
OTHER MATERIAL. Jones & Sherborn (1862 : 262) stated that Cytheridea penta-
gonalis was represented by a single specimen only. Two single right valves of this
species, 10.273 and 10.3929, also occur in the Winwood Collection from the Blue
Fuller's Earth Clay but cannot be regarded as paralectotypes.
DESCRIPTION. Carapace quadrate with sharply angled, prominent cardinal
angles and steeply sloping antero- and postero-dorsal slopes in the female dimorph,
elongate, more rectangular in outline in the male. Dorsal margin straight, ventral
margin medially incurved. Anterior end broadly rounded in the left valve, truncated
dorsally in the right by the steeply sloping antero-dorsal slope. In the queried
juvenile right valve this truncation of the anterior margin is not present. Posterior
end triangular, the postero-dorsal slope strongly concave in the female, resulting in
a distinct upturning of the posterior end in both valves but more strongly so in the
right. In the male this upturning is restricted to the right valve. Greatest length
of carapace below mid-point in both dimorphs but more ventrally situated in the
female. Greatest height and width developed in the posterior third of the carapace.
In the male the increase in width posteriorly is uniformly developed in that area. In
the female the increase in width tends to be restricted to the postero-ventral region.
Left valve larger than the right, with the left overlapping the right along the ventral
408 REVISION OF SOME
^^ tvv.'Vv'i-'.v.
.-.••.•:'::'X ttS.':: : •-•.••.•.•.•...'•;'<
m • MW
FIG. 7. Hadrocytheridea dolabra (Jones & Sherborn). Muscle scars of lectotype of
Cytheridea puteolata Jones & Sherborn, from the exterior. 1.1844. X 500.
margin. Elsewhere there does not appear to be any overlap. Shell surface very
finely punctate in between the large circular pits, each of which has a large normal
pore canal opening at its centre. Hinge antimerodont, strongly developed with five
anterior and five posterior teeth. Median groove loculate. Inner margin and line
of concrescence coincide to produce a broad duplicature through which pass (anteri-
orly) nine, long, straight and widely-spaced radial pore canals. Muscle scars
consist of a vertical row of four small, oval adductor scars, a small, round antero-
ventral mandibular scar and a large antero-dorsal antennal scar which is equal in
size to two adductor scars and is either heart-shaped or bill-hooked in outline.
DIMENSIONS. 1.1851, female right valve, length 0-64 mm. ; height 0-38 mm.
1.1844, male left valve, length 0-71 mm.; height 0-34 mm. 1.1859, male carapace,
length 0-73 mm. ; height 0-36 mm. ; width 0-37 mm. 1. 1866, female left valve,
length 0-57 mm. ; height 0-32 mm. 10.3294, male right valve, length 0-65 mm. ;
height 0-31 mm. 1. 1868, Pjuvenile right valve, length 0-46 mm. ; height 0-26 mm.
REMARKS. Jones & Sherborn (1888) described four different species which were
either male or female dimorphs of a single species. Although Cytheridea puteolata
had page preference it was decided to select a female dimorph as type because it is
the female outline which is continuous with juvenile instars. Of the two female
specimens available, that identified as Cytheridea dolabra was the better preserved
and was therefore chosen to identify the species. The breakdown of the synonymous
species is as follows:
Female right valve . . Cytheridea dolabra
Female left valve . . . Cytheridea pentagonalis
Male carapace . . . Cytheridea parattela
Male left valve . . . Cytheridea puteolata
Pjuvenile instar . . . Cytheridea ignobilis
ENGLISH BATHONIAN OSTRACODA 409
Hadrocytheridea dolabra is close to Cytheridea punctiputeolata Jones & Sherborn,
as far as the male right valve is concerned, but C. punctiputeolata has a lophodont
hinge, more rounded posterior end and a more strongly developed ornamentation of
punctae and pits. Cytheridea ignobilis has been included in this species as it probably
represents a juvenile instar. However, because of the slight difference between the
anterior margin of C. ignobilis and that of H . dolabra and the absence of other juveniles
to establish an ontogentetic sequence, C. ignobilis is doubtfully referred to this species
and is not placed in the list of paralectotypes.
Family PROGONOCYTHERIDAE Sylvester-Bradley 1948
Subfamily PROGONOCYTHERINAE Sylvester-Bradley 1948
Genus RECTOCYTHERE Malz 1958
Rectocythere sugillata (Jones & Sherborn)
(PL 10, figs. 3-6)
1888 Cytheridea sugillata Jones & Sherborn : 262, pi. 2, figs. -za-c.
DIAGNOSIS. Rectocythere with surface ornamentation of irregular ridges and raised
areas giving parched or shrivelled-up appearance. Coarsely pitted in well-preserved
specimens.
LECTOTYPE. 1.1855, lert valve figured Jones & Sherborn 1888. Blue Fuller's
Earth Clay, Midford.
PARALECTOTYPE. 10.3930, right valve referred to by Jones & Sherborn 1888 : 262,
Blue Fuller's Earth Clay, Win wood Collection.
DESCRIPTION. Shell robust, with high anterior and posteriorly sloping dorsal
margin. Anterior end broadly rounded; posterior end narrowly rounded in the left
valve, triangular in the right, with a concave postero-dorsal slope producing an
upturned posterior end. Ventral margin slopes upwards towards the posterior end,
but is over-hung postero-ventrally by the prominent convexity of the ventro-lateral
margin in that region. Anterior and posterior marginal borders flattened, the convex
central part of the carapace is strongly ornamented by irregular ridges and raised
areas which give a dehydrated appearance to the ostracod. Hinge lophodont.
Inner margin and line of concrescence coincide producing a rather broad duplicature.
To the outside of this there is developed a narrow flange extending around the
anterior margin and along the ventral margin to the apex of the posterior end.
Anterior radial pore canals long and straight, widely spaced and about seven to
eight in number. Muscle scars not observed.
DIMENSIONS. 1.1855, left valve, length 0-50 mm.; height 0-31 mm. 10.3930,
right valve, length 0-51 mm. ; height 0-32 mm.
REMARKS. Rectocythere sugillata possesses the same basic ornamentation as
R. rugosa Malz (19660 : 405, figs. 6-9) but may be distinguished by having a much
GEOL. 17, 8 25
4io REVISION OF SOME
finer degree of surface pitting and additional irregular swellings. Dr. Malz kindly
sent material of R. rugosa for comparison.
The ostracod Camptocythere lincolnensis Bate (1963 : 201, pi. 10, figs. 2-13) also
belongs to Rectocythere on its similarity of carapace outline, ornamentation and
internal details.
Genus CAYTONIDEA Bate 1965
DIAGNOSIS. Progonocy thermae, oval-rectangular in outline with well-rounded
anterior and posterior margins. Low eye swelling situated at anterior cardinal
angle. Cardinal angles prominent, broadly rounded. Shell surface with reticulate
ornamentation. Hinge antimerodont. Muscle scars consist of sub vertical row of
four adductor scars, rounded antero-dorsal antennal scar and rounded antero-ventral
mandibular scar. Radial pore canals long, straight, few in number. Left valve
larger than right. Dimorphic.
REMARKS. Since the publication of this genus a second species has been identified
which enables some modification to the diagnosis to be made. This concerns the
identification of dimorphism and the determination of a reticulate ornamentation as
a constant feature.
A re-examination of the type species, Caytonidea faveolata Bate (1965 : 100, pi. I,
figs. 13-14, pi. 2, figs, i-io), has shown that the holotype (10.1831) is a female dimorph
and the paratype (10.1834) a male.
Caytonidea terraefullonicae (Jones & Sherborn)
(PI. 10, figs. 2, 7, 8;P1. u, figs. 1-3)
1888 Cytheridea terrae-fullonicae Jones & Sherborn : 258, pi. i, figs. 5«-c.
DIAGNOSIS. Caytonidea with coarse reticulate ornamentation of five- to six-sided
pits covering entire shell surface. Extreme postero-ventral margin swollen, slightly
overhanging ventral surface. Males more elongate in outline than females.
LECTOTYPE. 1.1869, left valve, figured Jones & Sherborn 1888. Blue Fuller's
Earth Clay, Midford.
PARALECTOTYPES. 10.3917-20, male and female specimens from the Winwood
Collection, Blue and Yellow Fuller's Earth Clay, Midford.
OTHER MATERIAL. 10.4002-3, right valve and juvenile carapace from the J. F.
Blake Collection, Fuller's Earth Clay, Bath.
DESCRIPTION. Carapace rectangular in outline, especially in the male dimorph,
with sub-parallel dorsal and ventral margins. Anterior end high with greatest
height through the anterior cardinal angle which is situated close to the anterior
margin. Anterior end broadly but slightly obliquely rounded and extended ventrally
below the ventral margin. Posterior end broadly rounded in the left valve, with
ENGLISH BATHONIAN OSTRACODA 411
oblique postero-dorsal slope in the right valve. Postero- ventral margin slightly
swollen and overhanging ventral surface. Greatest length of carapace passes
slightly below the mid-point; greatest width in the posterior third. There are no
marginal borders. Left valve slightly larger than the right which it overlaps along
the ventral margin but very little elsewhere. Shell surface coarsely reticulate, the
reticulations producing five- to six-sided pits which cover the entire shell surface.
In the juvenile carapace, the reticulations tend to form longitudinal ridges along the
ventral surface. Normal pore canals large and widely spaced over the surface of
the carapace. Inner margin and line of concrescence coincide to produce a broad
duplicature through which pass a small number (about seven) of long, straight, and
widely spaced anterior radial pore canals. A narrow flange extends around the
anterior and ventral margins and along the postero-ventral part of the posterior
margin. Hinge antimerodont, the median element being long and finely denticulate/
locellate. Terminal elements dentate/loculate. Muscle scars as seen from the
exterior consist of a small, rounded antero-dorsal antennal scar and a vertical row of
four rounded adductor scars (type A). An oval smooth area in the region of the
anterior cardinal angle is indicative of an eye swelling. This feature is only clearly
seen in the male dimorph.
DIMENSIONS. 1.1869, female left valve, length 0-54 mm. ; height 0-30 mm.
10.3919, female right valve, length 0-49 mm. ; height 0-26 mm. 10.3917, male right
valve, length 0-54 mm. ; height 0-26 mm. 10.3918, male left valve, length 0-55 mm. ;
height 0-26 mm. 10.3920, female carapace, length 0-52 mm. ; height 0-29 mm. ;
width 0-25 mm.
REMARKS. Caytonidea terraefullonicae differs from C. faveolata Bate in the posses-
sion of a much stronger reticulate ornamentation, straighter dorsal margin, and
postero-ventral swelling slightly overlapping the ventral surface. Unfortunately,
during the examination of this material 10.3918 was lost and 10.3917 damaged.
Genus AC AN T HOC Y THERE Sylvester-Bradley 1956
Acanthocythere sphaerulata (Jones & Sherborn)
(PI. n, figs. 4-6)
1888 Cythere sphaerulata Jones & Sherborn : 253, pi. i, figs. 6a-c.
1956 Acanthocythere sphaerulata (Jones & Sherborn) Sylvester-Bradley : 12, pi. i, figs. 1-4.
DIAGNOSIS. As for Sylvester-Bradley 1956 : 12.
HOLOTYPE. 1.1835, female carapace, length 0-52 mm. ; height 0-31 mm. ; width
0-31 mm. Figured Jones & Sherborn 1888. Blue Fuller's Earth Clay, Midford.
REMARKS. Cythere sphaerulata was made the type species of Acanthocythere
(Sylvester-Bradley 1956), additional material indicating that the species was
dimorphic. A misprint on p. 12 stated that the holotype was a male dimorph ; it is,
in fact, a female dimorph.
4i2 REVISION OF SOME
Genus FASTIGATOCYTHERE Wienholz 1967
Fastigatocythere juglandica (Jones)
(PI- 12, fig. 3)
1888 Cythere juglandica var. major Jones & Sherborn : 255, pi. 4, figs. 2a, b.
REMARKS. The synonymy and diagnosis of this species are given on p. 389. The
variety described by Jones & Sherborn, of which the specimen 1.1872 is holotype
was correctly placed in synonymy by Sylvester-Bradley (1948).
DIMENSIONS. 1.1872, female left valve, length 0-74 mm. ; height 0-49 mm.
Genus GLYPTOCYTHERE Brand & Malz 1962
Glyptocythere guembeliana (Jones)
(PI. 3, fig. 2; PI. 4, fig. i)
1884 Cythere guembeliana Jones : 772, pi. 34, figs, 32, 33 [not fig. 31].
1888 Cytheridea pulvinar Jones & Sherborn : 266, pi. 3, figs. za-c.
1888 Cytheridea trapezoidalis Terquem, Jones & Sherborn : 270, pi. 4, figs. la, b.
1967 Glyptocythere guembeliana (Jones) Bate : 49, pi. 13, figs. 10-16, pi. 14, figs. 1-8.
REMARKS. This species has been described fully by me (Bate 1967) although
I omitted to include the ostracod identified by Jones & Sherborn as Cytheridea
trapezoidalis Terquem. This specimen is a juvenile instar of G. guembeliana.
Although it has a well-developed ventro-lateral keel, the outline of the valve is
typically juvenile in being strongly acuminate posteriorly. The hinge, in being anti-
merodont, reflects an early stage in the development of the adult entomodont hinge.
DIMENSIONS. 1.1858, female right valve (Cytheridea pulvinar}, length 0-82 mm. ;
height 0-47 mm. 1.1840, juvenile right valve (Cytheridea trapezoidalis}, length
0-54 mm. ; height 0-30 mm.
Glyptocythere oscillum (Jones & Sherborn)
(PI. 5, ng. 2; PI. 12, fig. 2; Text-figs. 8, 9)
1888 Cythere oscillum Jones & Sherborn : 254, pi. 3, figs. 8a-c.
1888 Cytheridea striblita Jones & Sherborn : 268, pi. 3, figs. ja-c.
DIAGNOSIS. Carapace strongly ornamented with series of grooves and irregular
swollen areas: two lateral grooves developed, the first ventro-laterally, the second
just above valve middle, both joined by narrow vertical groove passing between
two swollen areas. Broad dorso-median sulcus extends down to median groove.
Area below ventio-lateral groove alate in right valve, swollen in both valves. From
posterior region, two swollen areas extend towards valve centre whilst two similar
areas extend back from anterior region. Lower of anterior swollen area terminates
in prominent swelling. Anterior and posterior swollen areas do not unite. Normal
pore canals prominently displayed over carapace.
ENGLISH BATHONIAN OSTRACODA 433
HOLOTYPE. 1.1849, described, figured Jones & Sherborn (1888). Female right
valve from the Blue Fuller's Earth Clay, Midford.
OTHER MATERIAL. 10.3943, male left valve, referred to Jones & Sherborn (1888).
Winwood Collection. Blue Fuller's Earth Clay, Midford. This is the lectotype of
Cytheridea striblita Jones & Sherborn. 10.3914-6, two male left valves and a male
right valve, Winwood Collection, Blue Fuller's Earth Clay, Midford.
DESCRIPTION. Carapace subquadrate to sub-rectangular in outline, the more
elongate specimens being the males. Greatest length through the mid-point in the
male dimorph, below in the female right valve. Greatest height in the posterior
third. Dorsal margin, in the left valve, concave in the region of the dorso-median
sulcus, convex posteriorly. The dorsal margin of the right valve is strongly convex,
especially in the female. Cardinal angles particularly well developed in the right
valve, postero-dorsal slope concave in both valves, more especially so in the female
right valve. Antero-dorsal slope broadly convex in the male, concave in the female
right valve. Anterior end broadly rounded, posterior end triangular. Ventral
margin antero-medially concave, posteriorly convex. Ventro-lateral margin of
right valve becomes alate, extending below ventral surface. This feature does not
appear to be so well developed in the left valve. Anterior and posterior marginal
borders flattened. Shell surface ornamented as described in the diagnosis. Normal
pore canal openings prominent and widely scattered over the carapace. Hinge
with five to six terminal teeth in the right valve and a median loculate groove. In
the left valve the terminal loculate sockets are separated by a median bar which is
rather coarsely dentate, particularly in the anterior part. Although not as well
developed as in some species the hinge is regarded as entomodont. Inner margin
and line of concrescence coincide. Anterior radial pore canals long and straight,
widely spaced, about ten in number. Muscle scars not seen.
DIMENSIONS. Lectotype, 1.1849, female right valve, length 0-53 mm. ; height
0-30 mm. 10.3943, male left valve, length 0-62 mm. ; height 0-32 mm. 10.3916,
male right valve, length 0-59 mm. ; height 0-30 mm.
REMARKS. Jones & Sherborn identified the female specimens of the present species
as Cy there oscillum; the males they placed in Cytheridea striblita. The figured speci-
men of the latter species has been lost but the lectotype of C. striblita was found in
the Winwood Collection.
Glyptocythere rudimenta Brand & Malz (1962 and 1966) is similar to G. oscillum in
ornamentation but differs in that the anterior and posterior swollen areas take the
form of rather thin irregular ridges, not nearly so broad as in oscillum. A broad
sulcus is developed in oscillum in the dorso-median area, but in rudimenta there is a
series of irregular ridges radiating down from the dorsal margin. The shell outline
in the right valve is noticeably different, the cardinal angles and the dorso-median
extension of the dorsal margin being very much more prominently developed in
G. oscillum. The similarities in ornamentation between the two species suggests
a phylogenetic relationship which is supported by their stratigraphic position,
G. rudimenta being found in the Upper Bajocian and G. oscillum in the Upper
REVISION OF SOME
FIG.
Glyptocythere oscillum (Jones & Sherborn). Male left valve, paralectotype,
10.3943- x 153.
FIG. 9. Glyptocythere oscillum (Jones & Sherborn). Male right valve, 10.3916. x 165.
FIG. 10. Glyptocythere rudimenta Brand & Malz. Left valve, female carapace. 10.3921.
X 113.
ENGLISH BATHONIAN OSTRACODA 415
Bathonian. The kindness of Dr. D. E. Brand in sending me specimens of G. rudi-
menta is acknowledged here.
Glyptocythere persica (Jones & Sherborn)
(PL 12, fig. i; Text-fig, u)
1888 Cytheridea persica Jones & Sherborn : 270, pi. 4, fig. 4.
DIAGNOSIS. Carapace with broad dorso-median depression in anterior half; short,
blade-like ventral keel on ventral surface and ventro-lateral margin develops into
keel-like ridge. Lateral surface ornamented by neat reticulation. Ventral surface
with low, longitudinal ridges. Anterior and posterior marginal borders smooth,
compressed.
HOLOTYPE. 1.1834, right valve, figured Jones & Sherborn (1888), Blue Fuller's
Earth Clay, Midford.
OTHER MATERIAL. 10.4006, juvenile carapace, Mockler Collection, Fuller's Earth,
Midford.
DESCRIPTION. Carapace with greatest height at the anterior end, through the
anterior cardinal angle, tapering to the posterior end. Dorsal margin broadly
convex, sloping to the posterior, ventral margin with a broad, median incurvature,
below which in the right valve there is lip-like extension of the flange. The flange
is continuous around the anterior margin and along the ventral margin to the tip of
the triangular posterior end. Anterior end broadly rounded, posterior end with
FIG. u. Glyptocythere persica (Jones & Sherborn). Internal view of right valve.
Holotype, 1.1834. x I2°-
concave postero-dorsal slope and convex postero- ventral slope. Anterior and
posterior marginal borders compressed and smooth, distinct from the more strongly
convex part of the carapace which is ornamented by a neat reticulation. The
ornamentation is extended along the ventro-lateral margin to form a keel-like ridge.
Ventral surface ornamented by four longitudinal ridges, one of which is developed
416 REVISION OF SOME
into a short, blade-like keel. A smooth area at the anterior cardinal angle is sugges-
tive of an ocular lens. Just anterior of valve centre and in the dorso-median part
of the carapace a broad shallow sulcus is developed, especially noticeable in dorsal
view. The posterior part of the valve behind the sulcus is noticeably more convex,
whilst at the base of the sulcus, a raised area locates the position of the muscle scars.
The left valve is larger than the right, which it overlaps along the ventral margin and
slightly overreaches along the dorsal margin. The inner margin and line of con-
crescence coincide. The hinge is weakly entomodont.
DIMENSIONS. 1.1834, right valve, length 0-70 mm. ; height 0-38 mm. 10.4006,
juvenile carapace, length 0-52 mm. ; height 0-28 mm. ; width 0-24 mm.
REMARKS. Glyptocythere persica, with its neat, reticulate ornamentation, distinct
ventro-lateral ridge, and well-developed ventral, blade-like keel, is quite distinct
from those species which possess well-developed ventro-lateral ridges or keels, e.g.
Glyptocythere costata Bate (1965) and G. guembeliana (Jones) (Bate 1967), or from
those which have a much stronger reticulate ornamentation, e.g. Glyptocythere
tuberodentina Brand & Malz (19620).
Genus LOPHOCYTHERE Sylvester-Bradley 1948
BRADIANA GROUP
Lophocythere acutiplicata (Jones & Sherborn)
(PI. i, fig. 8; PI. 12, figs. 4, 6)
1888 Cytheridea acutiplicata Jones & Sherborn : 271, pi. 4, figs, 'ja-b, Sa-c.
DIAGNOSIS. Carapace dimorphic, tapering posteriorly. Eye swelling below
anterior cardinal angle. Shell surface weakly ornamented laterally, the most
prominent ridge being situated ventro-medially on the valve and extending from
posterior margin to anterior margin. A second lateral ridge just above also extends
down to the anterior margin, but cannot be traced backwards beyond the anterior
third where it dies out. Two short ridges branch off the ventro-median ridge a short
distance back from the anterior margin. The uppermost ridge is very short and
extends into valve centre at an acute angle whilst the lower extends below the ventro-
median ridge, parallel to it for almost three-quarters of its length. Three short
lateral ridges are situated in the dorso-median part of the posterior third. A short
ridge extends down from the eye swelling to die out above mid-length. Ventro-
median part of carapace convex, extended below ventral surface. On this extension
the left valve has a single ridge extending almost from the anterior margin to the
posterior margin, with a short lateral ridge below it in the anterior part of the valve.
In the right valve there are two lateral ridges in this region, both convex ventrally.
The upper extends the length of the posterior half of the valve whilst the ridge
slightly offset below it extends the length of the anterior half with some overlap of
the two at valve middle. Normal pore canals prominent. Hinge antimerodont.
Left valve larger than right.
ENGLISH BATHONIAN OSTRACODA 417
LECTOTYPE. 1.1863, female carapace, figured Jones & Sherborn (1888), Blue
Fuller's Earth Clay, Midford.
PARALECTOTYPES. 1.1847, female right valve, figured Jones & Sherborn
Blue Fuller's Earth Clay, Midford. 10.3642-45, two male and one female right valve
from the Blue Fuller's Earth Clay and one female right valve from the Yellow Fuller's
Earth Clay, Midford, Winwood Collection.
DESCRIPTION. Carapace with a strong posterior taper in both dimorphs, of which
the male is the more elongate. Anterior end high, broadly rounded, the greatest
height being through the anterior cardinal angle, just below which is a low eye
swelling. Posterior end triangular, narrow, with concave postero-dorsal slope and
convex postero- ventral slope. Posterior cardinal angle prominent. Greatest length
passes through the mid-point, greatest width in the posterior third although the
width in the anterior third is almost the same. Carapace constricted slightly about
centre when viewed dorsally. Normal pore canals large and prominently displayed
over the carapace. Shell surface ornamented as in the diagnosis. Left valve
larger than the right, which it overlaps along the ventral margin and overreaches
along the dorsal margin. Hinge antimerodont, only seen in the right valve
where there are four to five terminal teeth and a rather long, delicate, loculate median
groove. The hinge is particularly delicate, and the carapace as a whole appears to
be thin-shelled. Inner margin and line of concrescence coincide. Radial pore
canals and muscle scars not observed. A narrow flange extends around the
anterior and ventral margins and along the ventral part of the posterior.
DIMENSIONS. 1.1863, female carapace, length 0-57 mm.; height 0.31 mm.; width
0-25 mm. 1.1847, female right valve, length 0-54 mm. (broken); height 0-32 mm.
10.3643, male right valve, length 0-65 mm. ; height 0-33 mm. 10.3644, female right
valve, length 0-58 mm. ; height 0-33 mm. 10.3645, female right valve, length
0-56 mm. ; height 0-31 mm.
REMARKS. The carapace morphology and surface ornamentation agree closely
with the basic characters of those ostracods which may be placed in the bradiana
grouping of Lophocy 'there. Apart from the details of the radial pore canals and
muscle scars, which are not so far known for L. acutiplicata, the only character in
disagreement with this classification is the hinge, which, being antimerodont, is at
once at variance with all the other species of Lophocythere. An antimerodont hinge
is normally accepted as present in juvenile instars of adults having an entomodont
hinge; here the delicate shell, strong posterior taper and comparatively small size
would indicate this if it were not for the presence of strong dimorphism suggesting
adult status. Sexual dimorphism is known, however, in pre-adult instars. As only
right valves are available for the examination of internal features, it is not intended
to suggest a subgeneric rank for this species, until additional material becomes
available. In ornamentation L. acutiplicata may be distinguished easily by the
number of lateral ridges, their tendency to fade out in the posterior part of the
carapace, and by the development of two ventro-lateral keels in the right valve.
418 REVISION OF SOME
Lophocythere bradiana (Jones)
(PL 4, figs. 2, 3; Text-fig. 4)
REMARKS. The species has already been described (p. 390).
DIMENSIONS. 1.1854 (lectotype of C. craticula figured Jones & Sherborn, pi. 4,
figs. loa-c), length 0-62 mm.; height 0-32 mm.; width 0-32 mm. 1.1867 (paralecto-
type of C. craticula figured Jones & Sherborn, pi. 4, figs, ga-c) left valve, length
0-60 mm.; height 0-32 mm.
Lophocythere septicostata Bate
(PI. 12, fig. 8)
1888 Cytheridea bradiana Jones & Sherborn : 272, pi. 4, figs. na-c.
1967 Lophocythere septicostata Bate : 52, pi. 15, figs. 7-13, pi. 16, figs. 1-4.
DIAGNOSIS. Lophocythere having seven longitudinal ridges on lateral surface and
two on ventral surface. Vertical ridge extends downwards from oval eye swelling
Interspaces between ridges punctate, with large, circular, normal pore canal openings.
Anterior and posterior marginal borders compressed.
REMARKS. For remarks relating to this species see Bate (1967).
DIMENSIONS. 1.1843, female right valve, length 0-64 mm.; height 0-33 mm.
Lophocythere sp.
(Text-fig. 12)
REMARKS. A single left valve in the Winwood Collection represents a new species
of Lophocythere. It comes from the Blue Fuller's Earth Clay of Midford and is of
the Bradiana Group. The arrangement of the lateral ridges is quite different from
that in any described species. The dorsal ridge commences at the anterior margin,
bends sharply upwards to pass through the eye swelling and then in an undulating
FIG. 12. Lophocythere sp. External view, right valve. 10.3900. x 170.
ENGLISH BATHONIAN OSTRACODA 419
course parallels the dorsal and postero-dorsal margins to die out at the extreme
posterior. The median ridge commences at the anterior margin just below the dorsal
ridge and soon bifurcates, the ventral branch extending back into the posterior
third. A second ridge branches from the median ridge a short distance above the
first bifurcation and extends into the posterior third where it unites with the first
bifurcation. Both then extend to the posterior margin as a single narrow ridge.
The median ridge continues its dorsal course and bifurcates yet again, the dorsal
branch continuing until it fuses with the dorsal ridge just behind the eye swelling;
the other branch passes laterally back into the posterior half of the valve, splits into
a reticulate complex of smaller ridges but re-forms to fuse with the dorsal ridge in
the posterior quarter. Two ventral ridges originate at the anterior margin, the
lower extending back into the posterior half of the valve whilst the upper is restricted
to the anterior half only. Interspaces between the ridges are reticulate. Hinge
strongly entomodont.
DIMENSIONS. 10.3900, left valve, length 0-50 mm. ; height 0-27 mm.
OSTREATA GROUP
Lophocy there fulgurata (Jones & Sherborn)
(PL 12, fig. 7)
1888 Cytheridea fulgurata Jones & Sherborn : 273, pi. 4, figs. i2a-c.
DIAGNOSIS. Carapace sub-rectangular, tapering to the posterior. Oval, clear
eye swelling at anterior cardinal angle. Anterior and posterior marginal areas
flattened, smooth. Central part of carapace strongly convex, ornamented by two
prominent ridges. The first, L-shaped, commences at the eye swelling, extends
vertically down to the an tero- ventral part of the valve, where it performs a right-
angle, and runs laterally backwards to the edge of the posterior marginal area.
A second lateral ridge arises just below the right-angled bend of the first ridge and
extends backwards, paralleling the ridge above it, to die out in a similar position at
the edge of the posterior marginal border. The area inside the " L " of the first
ridge is occupied by strong ridges radiating down from the dorsal margin and
occasionally branching.
LECTOTYPE. 1.1832, right valve, figured Jones & Sherborn (1888), Blue Fuller's
Earth Clay, Midford.
PARALECTOTYPES. 10.3646-8, right and left valves from the Blue Fuller's Earth
Clay, Midford, Winwood Collection.
DESCRIPTION. Carapace sub-rectangular with dorsal margin sloping to posterior.
Cardinal angles prominent, situated close to the anterior and posterior margins
respectively. Anterior cardinal angle smooth, indicating the presence of an eye
swelling. Line of greatest height passes through this point. Line of greatest
length passes through the mid-point, or slightly below it. Anterior end high,
420 REVISION OF SOME
broadly rounded; posterior end small and triangular with a concave postero-dorsal
slope and a convex postero-ventral slope. Ventral margin with a shallow antero-
median incurvature. Shell surface ornamented as in the diagnosis; area between
the ridges appears to be punctate. Hinge not well preserved, the median element in
both valves having been damaged. Terminal elements consist of dentate ridges in
the right valve and loculate sockets in the left. Median bar of the left valve, long,
not clearly seen to be dentate, bounded above by a shallow accommodation groove.
Median groove of the right valve, long and narrow, not seen to be loculate because of
damage. Inner margin and line of concrescence coincide. Radial pore canals and
muscle scars not seen. A narrow flange extends around the anterior margin and
along the ventral margin to the tip of the posterior margin in the right valve.
DIMENSIONS. 1.1832, right valve, length 0-77 mm. ; height 0-42 mm. 10.3648,
left valve, length 0-66 mm. ; height 0-36 mm.
REMARKS. Although the median element of the hinge has been damaged, the
appearance suggests that it was antimerodont instead of the entomodont hinge
typical of the genus. The strongly tapering outline and the hinge suggest that the
material is of a pre-adult instar, but it is doubtful whether the adult ornamentation
would be different from that described above. Certainly there is no difference
between the instars examined.
Lophocythere ostreata (Jones & Sherborn)
(PI. 12, fig. 5; Text-fig. 13)
1888 Cytheridea ostreata Jones & Sherborn : 271, pi. 4, figs. 6a-c.
1888 Cytheridea bicarinata Jones & Sherborn : 270, pi. 4, figs. 50— c.
1948 Lophocythere ostreata (Jones & Sherborn) Sylvester-Bradley : 195, pi. 14, figs. 1—4, pi. 15,
figs, i, 2.
1963 Lophocythere ostreata (Jones & Sherborn); Oertli : 42, pi. 28, fig. o, pi. 29, fig. o.
DIAGNOSIS. Species of Lophocythere with characteristic L-shaped ridge com-
mencing at a large, clear eye node, extending down to the antero-ventral region, to
bend back at a right-angle and extend to the edge of the posterior marginal border.
Second, keel-like ridge starts below the previous ridge in the antero-ventral part of
the valve and extends backwards, parallel to the first ridge to die out against the
edge of the posterior marginal border. Shell surface coarsely reticulate, the reticula-
tions strongly upstanding to produce an oblique row of three prominent areas, the
first of which is antero-central in position, just inside the angle of the L-shaped
ridge. The second is dorso-median in position, just behind mid-point, whilst the
third is postero-dorsal in position. Two, sometimes three, minor raised reticulations
may be situated below and behind the two most dorsal of the major raised areas.
Ventral surface with minor longitudinal ridges.
HOLOTYPE. 1.1833, right valve figured Jones & Sherborn (1888), Yellow Fuller's
Earth Clay, Midford.
ENGLISH BATHONIAN OSTRACODA 421
OTHER MATERIAL. 10.3945-7, two carapaces and two broken right valve frag-
ments, Winwood Collection, Blue Fuller's Earth Clay, Midford. These were referred
to by Jones & Sherborn (1888) and are syntypes of Cytheridea bicarinata, the figured
specimen having been lost. 10.3945 is here selected lectotype of C. bicarinata.
FIG. 13. Lophocythere ostreata (Jones & Sherborn). Internal view, right valve.
Holotype, 1.1833. X IO5-
DESCRIPTION. Carapace sub-rectangular with sub-parallel dorsal and ventral
margins in the adult. Juvenile specimens have distinct posterior taper. Anterior
end high, broadly rounded, posterior end triangular, with concave postero-dorsal
slope and convex postero-ventral slope. Cardinal angles prominent, rounded.
Anterior cardinal angle with well-developed, oval, clear eye node situated just below.
Line of greatest length passes through the mid-point, line of greatest height through
anterior cardinal angle and of greatest width through the posterior third of the
carapace. Ornamentation as in the diagnosis. The reticulations continue into the
lateral ridges and pass on to the anterior and posterior marginal borders. The
postero-dorsal raised area is not developed in juvenile instars. Left valve larger than
the right, which it overlaps along the ventral margin but overreaches around the
anterior and posterior margins and in the region of the cardinal angles. Hinge
weakly entomodont, seen only in the right valve: anterior dentate ridge with six
teeth, posterior element not seen, median groove poorly preserved and not definitely
seen to be loculate, expanded slightly in the anterior half. Inner margin and line
of concrescence coincide to produce a broad duplicature. Selvage distinct, flange
developed around the anterior margin, extending from the anterior cardinal angle,
along the ventral margin to the tip of the posterior margin. Radial pore canals
and muscle scars not observed.
DIMENSIONS. 1.1833, right valve, length 0-81 mm. ; height 0-40 mm. 10.3945,
carapace, length 0-85 mm. ; height 0-43 mm. ; width 0-42 mm. 10.3947, juvenile
carapace, length 0-60 mm. ; height 0-31 mm. ; width 0-28 mm.
REMARKS. The Ostreata Group of Lophocythere is characterized by the presence
of two lateral keel-like ridges, the upper of which turns upwards to produce a reverse
L-shaped ridge. The ornamentation within this " L " varies according to the
species. In L. ostreata the reticulate ornamentation, extended into a number of
upstanding areas, is quite unlike any described species. Exaggerations in the
ornamentation of species within this group tend to be in the same position as the
areas emphasized in L. ostreata. In this connection the closest species is L. scabra
Triebel (1951), although it does not possess the strong reticulate ornamentation of
ostreata.
422 REVISION OF SOME
Genus MICROPNEUMATOCYTHERE Bate 1963
Micropneumatocythere limaciformis (Jones & Sherborn)
(PI. 13, fig. i)
1888 Cytheridea limaciformis Jones & Sherborn : 269 pi. 3 figs. i2a-c.
DIAGNOSIS. Carapace oval/elongate with dorso-median area swollen, convex,
projecting above dorsal margin in right valve; left valve with convex dorsal margin.
Ventro-lateral and median part of ventral surface swollen, strongly convex, some-
times with weak longitudinal striae on ventral surface of convexity. Shell surface
punctate. Normal pore canal openings laige, circular, evenly spaced over the
carapace.
HOLOTYPE. 1.1831, male right valve, figured Jones & Sherborn (1888), Blue
Fuller's Earth Clay, Midford.
OTHER MATERIAL. 10.276 and 4007, and 10.3944, female right and left valves
from the Bradford Clay, Bradford-on-Avon, and male right valve from the Blue
Fuller's Earth Clay, Midford. Win wood Collection.
DESCRIPTION. Carapace oval/elongate in outline, the more elongate specimens
being the males. In the right valve of both dimorphs, especially the male, the
dorso-median part of the valve is distinctly swollen, the convex surface projecting
above the dorsal margin. Similarly the ventro-lateral part of the valve is also
swollen but involves the ventral surface where, in some specimens, weak longitudinal
striae are present. Left valve with uniformly convex dorsal and ventral margins.
Anterior end unifoimly rounded, posterior end triangular with concave postero-dorsal
slope and convex postero-ventral slope, the posterior extremity tending to be
upturned. Cardinal angles particularly distinct in the right valve, apparently
separated from the central convexity of the valve by a shallow groove. Greatest
length of carapace passes through the mid-point with the greatest height and width
also at valve centie. Shell surface finely punctate with prominent, large, circular
normal pore canal openings evenly spaced over the carapace. Hinge almost certainly
antimerodont, the terminal dentate/loculate elements being clearly seen, whilst the
median bar and groove is not sufficiently well preserved to state that it is a dentate/
loculate structure. Inner margin and line of concrescence coincide, anterior radial
pore canals straight, evenly spaced around the anterior margin, exact number not
observed but no more than eight. A narrow flange extends around the anterior
margin of the right valve and along the median part of the ventral margin. Muscle
scars not seen.
DIMENSIONS. 1.1831, male right valve, length 0-71 mm. ; height 0-37 mm.
10.3944, male right valve, length 0-62 mm. ; height 0-32 mm. 10.276, female right
valve, length 0-56 mm. ; height 0-36 mm. 10.4007, female left valve, length
(damaged) 0-61 mm. ; height 0-37 mm.
REMARKS. The closest species is M. quadmta Bate (1967), particularly in the case
of the male dimorph. However, the characteristic development of a deep postero-
ENGLISH BATHONIAN OSTRACODA 423
ventral slope in M. quadrata and the less strongly developed dorso-median convexity
of the right valve distinguish this species from M. limaciformis.
Genus PROGONOCYTHERE Sylvester-Bradley 1948
Progonocythere stilla Sylvester-Bradley
(PL 13, ng. 2)
1948 Progonocythere stilla Sylvester-Bradley : 190, pi. 12, figs, i, 2, pi. 13, figs, i, 2.
1956 Progonocythere stilla Sylvester-Bradley; Grekoff : pi. 13, figs. 325-7.
1963 Progonocythere stilla Sylvester- Bradley ; Oertli : 44, pi. 28, fig. n, pi. 29, fig. n.
DIAGNOSIS. Progonocythere with finely punctate shell surface, large, circular
normal pore canal openings, somewhat irregularly scattered, tending to be con-
centrated towards centre of valve. Anterior end high, broadly rounded. Anterior
cardinal angle close to anterior margin. Cardinal angles well rounded in left valve
more distinctly angled in right. Dorsal margin convex in right valve, straight in left,
in both valves steeply sloping to posterior. Posterior rounded in left valve, slightly
angled in right. Anterior marginal border distinct from convexity of valve, ventro-
lateral margin strongly convex, overhanging ventral margin. Left valve larger
than the right. Hinge entomodont. Other internal features as for genus.
Dimorphic.
REMARKS. Progonocythere stilla, the type species ol the genus, was described by
Sylvester-Bradley from the Bathonian Boueti Bed of Langton Herring. A single
left valve (10.3657) found in the Winwood Collection, from the Blue Fuller's Earth
Clay of Midford, was not described by Jones & Sherborn. Three specimens
(10.3584-6) in the Mockler Collection are from the Fuller's Earth of Midford.
DIMENSIONS. 10.3657, left valve, length 0-60 mm.; height 0-43 mm.
Genus TERQUEMULA Blaszyk & Malz 1965
Terquemula blakeana (Jones)
(PL 4, fig. 8)
REMARKS. For synonymy and diagnosis see p. 393. The two specimens in the
Jones & Sherborn Collection which have been placed in the synonymy are : Cytheridea
transversiplicata, figured Jones & Sherborn 1888, pi. 3, figs, ^a-c, 1.1828 (holotype),
right valve, length 0-59 mm.; height 0-31 mm., and Cytheridea blakeana (Jones)
Jones & Sherborn, figured pi. 2, figs. na-c. 1.1853, left valve, length 0-60 mm. ;
height 0-31 mm. In addition a single right valve, 10.3941, has been found in the
Winwood Collection from the Blue Fuller's Earth Clay, Midford.
INCERTAE SEDIS
The following ostracod species described by Jones & Sherborn are represented by
single specimens only. They are retained here under their original names until
comparative material enables a more precise determination.
424 REVISION OF SOME
" Cythere " corrosa Jones & Sherborn
(PL 13, fig- 4)
1888 Cythere corrosa Jones & Sherborn : 254, pi. 2, figs. 120— c.
HOLOTYPE. 1.1865, figured Jones & Sherborn (1888), Blue Fuller's Earth Clay,
Midford.
REMARKS. This small ostracod resembles Caytonidea in outline and reticulate
ornamentation, but differs in possessing a well-defined anterior marginal border.
In Caytonidea the convex outline of the carapace continues without a break on to the
anterior margin. Internal features not seen.
DIMENSIONS. 1.1865, carapace, length 0-49 mm.; height 0-28 mm.; width
0-28 mm.
" Cytheridea " aequabilis Jones & Sherborn
(PI. 13, fig. 3)
1888 Cytheridea aequabilis Jones & Sherborn : 257, pi. i, figs. la-c.
HOLOTYPE. 1.1864, figured Jones & Sherborn (1888), Yellow Fuller's Earth Clay,
Midford.
REMARKS. C. aequabilis has a finely punctate carapace, rounded anterior and
posterior margins, and parallel dorsal and ventral margins. In outline it is close to
the unidentified ostracod figured by Bate (1964, pi. 13, figs. 5-8). Internal features
not seen.
DIMENSIONS. 1.1864, carapace, length 0-71 mm.; height 0-30 mm ; width
0-32 mm.
" Cytheridea " coarctata Jones & Sherborn
(PL 13, fig- 5)
1888 Cytheridea coarctata Jones & Sherborn : 259, pi. i, figs. 3«-e.
HOLOTYPE. 1.1841, figured Jones & Sherborn (1888), Blue Fuller's Earth Clay,
Midford.
REMARKS. The unornamented carapace, well-rounded anterior and posterior
margins, parallel dorsal and ventral margins, together with a muscle scar pattern in
which the antero-dorsal antennal scar is crescentic in shape (type B), suggest
Homocytheridea Bate (i963«). The left valve, the only known specimen, has been
broken posteriorly and a more definite identification is not attempted.
DIMENSIONS. 1.1841, left valve, length (broken) 0-56 mm.; height 0-26 mm.
ENGLISH BATHONIAN OSTRACODA
425
" Cytheridea " eminula Jones & Sherborn
(PI. 13, fig. 6)
1888 Cytheridea eminula Jones & Sherborn : 261, pi. i, figs. na-c.
LECTOTYPE. 1.1839, figured Jones & Sherborn (1888), Blue Fuller's Earth Clay,
Midford.
REMARKS. This species is represented by a single left valve, although two speci-
mens were included in the original description.
DIMENSIONS. 1.1839, kft valve, length 0-54 mm. ; height 0-31 mm.
" Cytheridea " punctiputeolata Jones and Sherborn
(PI. 13, fig. 8)
1888 Cytheridea punctiputeolata Jones & Sherborn : 258, pi. i, figs.
HOLOTYPE. 1.1856, figured Jones & Sherborn (1888), Blue Fuller's Earth Clay,
Midford.
REMARKS. Although at first appearing to resemble the male dimorph of Hadro-
cytheridea dolabra the present species differs in the much more rounded posterior
margin, in the greater exaggeration of the ornamentation, which here consists of large,
circular pits with the interspaces between distinctly punctate, and in the lophodont
hinge.
DIMENSIONS. 1.1856, right valve, length 0-73 mm.; height 0-32 mm.
" Cytheridea " retorrida Jones & Sherborn
(PI. 15, fig. 3; Text-figs. 14, 15)
1888 Cytheridea retorrida Jones & Sherborn : 260, pi. i, figs. 8a-c.
MATERIAL. 10.3923, Blue Fuller's Earth Clay, Midford. Winwood Collection.
REMARKS. The figured specimen was never presented and is presumed lost.
A single carapace of this species has been found in the Winwood Collection, but
FIG. 14. Cytheridea retorrida Jones & Sherborn. Right valve, complete carapace.
10.3923. x 135.
26
GEOL. 17,
426 REVISION OF SOME
cannot be assigned to a known genus. Like C. corrosa it has some affinities with
Caytonidea but has been excluded on the presence of distinct marginal borders. No
internal features are known.
FIG. 15. Cytheridea retorrida Jones & Sherborn. Dorsal view of carapace with
ornamentation omitted. 10.3923. x 135.
DIMENSIONS. 10.3923, carapace, length 0-54 mm. ; height 0-28 mm.; width
0-28 mm.
" Cytheridea " spinifastigiata Jones & Sherborn
(PI. 13, ng. 7)
1888 Cytheridea spinifastigiata Jones & Sherborn : 264, pi. 2, figs. 8a-c.
HOLOTYPE. I.i86i, figured Jones & Sherborn (1888), Blue Fuller's Earth Clay,
Midford.
REMARKS. It is not possible to assign the single left valve to any known genus.
DIMENSIONS. 1. 1861, left valve, length 0-60 mm.; height 0-33 mm.
" Cytheridea " spinigyrata Jones & Sherborn
(PL 13, fig. 9)
1888 Cytheridea spinigyrata Jones & Sherborn : 264, pi. 2, figs. ja-c.
HOLOTYPE. 1. 1860, figured Jones & Sherborn (1888), Blue Fuller's Earth Clay,
Midford.
ENGLISH BATHONIAN OSTRACODA 427
REMARKS. Jones & Sherborn (p. 264) compared this species with Cythere drupacea
Jones but at the same time pointed out differences. The superficial similarity of
C. spinigyrata to C. drupacaea suggests that it, too, might belong to Metacytheropteron
Oertli (1957).
DIMENSIONS. 1. 1860, right valve, length 0-61 mm.; height 0-34 mm.
IV. THE J. F. BLAKE COLLECTION
Order MYODOCOPIDA Sars 1866
Suborder CLADOCOPINA Sars 1860
Family POLYCOPIDAE Sars 1866
Genus POLYCOPE Sars 1866
Polycope fungosa sp. nov.
(PI. 16, figs. 4, 9)
DIAGNOSIS. Carapace oval in adults, almost circular in juvenile instars. Shell
surface strongly pitted with weak striae near and parallel to the free margin.
Anterior end flattened, with marginal denticles. Hinge short, sunk in dorsal groove.
Anterior cardinal angle prominent, antero-dorsal slope long, broadly convex with
prominent overlap of left valve by the right. Posterior end broadly rounded, ventral
margin strongly convex. Carapace distinctly angled at junction of antero-dorsal
slope and anterior margin. Right valve larger than left.
HOLOTYPE. 10.3951, carapace, Fuller's Earth, Midford, near Bath.
PARATYPES. 10.3952-5, one adult and eighteen juvenile carapaces. Horizon
and locality as above.
DESCRIPTION. Carapace in adult specimens oval in outline, laterally compressed.
Lines of greatest length and height pass through the mid-point. Greatest width in
the holotype situated at the extreme posterior end. A shallow central depression
on the lateral surface marks the position of the muscle scars. Hinge margin situated
in the posterior half of the carapace and sunk into a dorsal groove, at the anterior
end of which the cardinal angle is prominently developed. Antero-dorsal slope long
and convex, terminating in a sharply angled junction with the obliquely flattened
anterior end. Posterior end broadly rounded, ventral margin broadly convex.
Shell surface uniformly and distinctly pitted; striae are developed in the anterior
and ventro-lateral regions. In dorsal view one stria on each valve can be seen to
project beyond the anterior margin. Striations are arranged parallel to the free
margins. Anterior margin may develop small denticles, though these, like the
surface ornamentation, are generally absent from the juvenile instars. Right valve
larger than the left, which it overlaps strongly along the antero-dorsal slope and less
strongly around the posterior end. Internal features not observed.
DIMENSIONS. 10.3951, carapace, length 0-54 mm. ; height 0-44 mm.; width
0-23 mm. 10.3952, carapace, length 0-46 mm.; height 0-37 mm.; width 0-20 mm.
4z8 REVISION OF SOME
10.3953, juvenile carapace, length 0-32 mm. ; height 0-27 mm. ; width 0-09 mm.
10.3954, juvenile carapace, length 0-26 mm. ; height 0-22 mm. ; width o-oi mm.
REMARKS. A number of species of Poly cope have been described from the Lower
Jurassic (Apostolescu 1959; Fischer 1961 and Blake 1876), Middle Jurassic (Terquem
1885) and Upper Jurassic (Donze 1962 and Oertli 1959). P. fungosa differs from
them all in the type of ornamentation present. A number of specimens in the Blake
collection labelled Poly cope sp. are probably juvenile specimens of the present species
but lack the specific ornamentation.
Family PROGONOCYTHERIDAE Sylvester-Bradley 1948
Genus TRACHYCYTHERE Triebel & Klingler 1959
REMARKS. Trachycythere Triebel & Klingler (1959 : 343), first described from the
Lias of Germany and later recorded from the Upper Bathonian of France (Oertli
1963), was placed in the Family Trachyleberididae (Moore 1961 : 341). Later Bate
(19670 : 551) transferred it to the Progonocytheridae, a classification retained in
this paper.
Trachycythere sp.
(PL 16, fig. 3)
REMARKS. A single left valve represents this Upper Bathonian species, which
differs in being sub-rectangular with almost parallel dorsal and ventral margins,
from two of the three species originally described by Triebel & Klingler, which have
a more pronounced posterior taper. Trachycythere verrucosa Triebel & Klingler
(1959) resembles this species in outline, but has stronger, more massive tubercles.
DIMENSIONS. 10.3966, left valve, length 0-45 mm. ; height 0-25 mm.
Genus HEKISTOCY THERE nov.
DERIVATION OF NAME. Hekistos (Gr.), least, + cythere.
DIAGNOSIS. A small genus with oval, posteriorly tapering carapace. Greatest
height close to the anterior margin. Left valve larger than right. Hinge lophodont.
Muscle scars of type A although antennal scar tends to be almost centrally situated
in front of adductors. Radial pore canals straight, widely spaced, ten anteriorly,
four posteriorly. Duplicature narrow.
TYPE SPECIES. Hekistocy there venosa sp. nov.
Hekistocy there venosa gen. et sp. nov.
(PI. 14, figs. 2, 4, 9; PI. 15, fig. 7; Text-figs. 16-18)
DIAGNOSIS. Hekistocy there with strongly ornamented carapace. Broad diagonal
ridge extends from postero-dorsal to antero- ventral region of valve. Short, thick,
ENGLISH BATHONIAN OSTRACODA
429
subsidiary ridges given off on dorsal and ventral sides of main ridge. Smaller
intermediary ridges producing a reticulate ornamentation. Diagonal ridge produced
at termination of, and at right-angles to, main ridge in postero-dorsal region.
HOLOTYPE. 10.3995, carapace, Fuller's Earth, Bath.
PARATYPES. 10.3996-9 (seven specimens) and 10.4011-3, Fuller's Earth, Bath.
DESCRIPTION. Carapace small with high, rounded anterior end and narrow,
rounded posterior end. Carapace parallel-sided in dorsal view. Line of greatest
length passes through the mid-point. Greatest width in either the anterior or
18
FIGS. 16-18. Hekistocy there venosa sp. nov. FIG. 16. Anterior radial pore canals. Para-
type, 10.4013. x 300. FIG. 17. Hinge, right valve, paratype, 10.4011. x 300.
FIG. 18. Muscle scars, left valve, paratype, 10.3997. X 300.
posterior third, the carapace being slightly constricted medially. Dorsal margin
convex, especially in the right valve where it projects slightly above that of the left
valve. Surface ornamentation very coarsely reticulate, produced by a series of
irregular ridges, the major of which extends down from the postero-dorsal part of the
valve and extends to the antero- ventral region. From this ridge a number of
irregular and shorter ridges branch off on its dorsal and ventral sides. One of the
secondary ridges is particularly noticeable in dorsal view, and this leaves the primary
ridge in the posterior part of the valve to extend dorsally in an antero-dorsal direction
until it fuses with the dorsal margin. Left valve larger than the right, which it
overlaps along the ventral margin and overreaches in the region of the antero- and
430 REVISION OF SOME
postero-dorsal slopes. Ventral surface flattened, triangular in outline, widening
towards the posterior end. Hinge lophodont: right valve with smooth terminal
elements which are the dorsal terminations of the selvage; median groove deeply
recessed, smooth. Left valve with smooth terminal sockets open ventrally to the
interior of the valve. Median bar long, smooth, convex, the convexity of the bar
facing upwards. Inner margin and line of concrescence coincide, the duplicature
being rather narrow. Radial pore canals short and straight and widely spaced,
ten anteriorly and four posteriorly. Muscle scars of type A (Bate 1963), with four
adductor scars in a slightly crescentic row; the second dorsal adductor is laterally
elongate with the rounded antennal scar situated in front.
DIMENSIONS. 10.3995, carapace, length 0-32 mm. ; height 0-20 mm. ; width
0-15 mm. 10.3996, left valve, length 0-32 mm. ; height 0-21 mm. 10.4011, right
valve, length 0-32 mm. ; height 0-20 mm. 10.4012, left valve, length 0-33 mm. ;
height 0-21 mm.
REMARKS. Hekistocy there has a distinctive combination of lophodont hinge, small,
oval, posteriorly tapered carapace, type A muscle scars, and simple radial pore
canals.
Family PROTOCYTHERIDAE Ljubimova 1955
Subfamily KIRTONELLINAE Bate 1963
Genus EKTYPHOCY THERE Bate 1963
Ektyphocythere parva (Oertli)
(PI. 14, figs, i, 3)
J959 Procytheridea minuta Oertli : 122, pi. 3, figs. 37-40.
1960 Procytheridea parva Oertli : 70.
REMARKS. Two complete carapaces represent this species, one male the other
female.
When first erected, Ektyphocythere Bate (1963) included those species of " Pro-
cytheridea " which possessed, amongst other characters, a triangular arrangement of
ridges as ornamentation and an antimerodont hinge. P. parva was one of the species
included.
Subsequently Malz (1966) placed Ektyphocythere in synonymy with Kinkelinella
Martin (1960) the type species, K. tenuicostati Martin (1960), having a reticulate
ornamentation. Although considered here to be closely related the clear distinction
between the forms having a triangular rather than a reticulate ornamentation is
such that the retention of the former in Ektyphocythere is maintained here.
DIMENSIONS. 10.3982, female carapace, length 0-41 mm. ; height 0-23 mm. ;
width 0-21 mm. 10.3983, male carapace, length 0-45 mm.; height 0-23 mm.; width
0-24 mm.
ENGLISH BATHONIAN OSTRACODA 431
FAMILY UNCERTAIN
Genus PLATYCYTHERE Bate 1967
Platycythere sp.
(PI. 14, fig. 5)
REMARKS. Represented by a single specimen only. In outline and possession
of dorsal and ventro-lateral swellings this species exhibits the generic features
described (Bate 1967), but differs from Platycythere verriculata, the type species, by
having a much finer reticulate ornamentation and a median swelling on the lateral
surface of both valves. This swelling, positioned about the centre of the valve, has
an oblique postero-dorsal antero- ventral inclination.
Gesoriacula Oertli (1959) has a similar external morphology but a more primitive
hinge structure.
DIMENSIONS. 10.3992, carapace, length 0-35 mm. ; height 0-20 mm. ; width
0-16 mm.
Family CYTHERURIDAE Miiller 1894
Genus LOONEYELLA Peck 1951
REMARKS. The type species, Cythere monticula Jones (1893), was described from
the Cretaceous Bear River Formation, Cokeville, south-west Wyoming, in association
with fresh- to brackish-water ostracods. Peck (1951), in resampling this Formation,
found a new ostracod species which he considered to be congeneric with C. monticula,
and for these two species erected the genus Looneyella, C. monticula being the type.
The preservation of Peck's material is considerably better than that of the holotype,
which is in the Museum collections. The latter, 1.5872, figured Jones (1893, pi. 15,
fig. 13), is a complete carapace, length 0-65 mm.; height 0-40 mm.; width 0-39 mm.
(inclusive of spines). The holotype is figured here (PL 14, fig. 6) for comparison with
the Bathonian species L. subtilis.
To date the following species have been placed in Looneyella :
L. monticula (Jones) from the Upper Cretaceous Bear River Formation.
L. quadrispina Peck 1951, also from the Bear River Formation.
L. subtilis Oertli, from the Bathonian of the Boulonnais, France.
Looneyella subtilis Oertli
(PL 14, figs. 7, 8)
1959 Looneyella ? subtilis Oertli : 119, pi. 3, figs. 31-5.
DIAGNOSIS. Small species of Looneyella with prominent postero-dorsal tubercle,
an antero-median tubercle somewhat smaller in size, and a marginal ridge which
extends around the anterior margin and along the ventro-lateral alar extension.
Both tubercles and marginal ridge are hollow when viewed internally. Antero-
432 REVISION OF SOME
medially there is a break in the continuity of the marginal ridge. Small swelling
may be present below the postero-dorsal tubercle, and at the termination of the alar
ridge. Shell surface smooth except for ventral surface which has four longitudinal
striae per valve. Left valve larger than right. Hinge antimerodont. Species
dimorphic. Eight straight anterior radial pore canals; inner margin and line of
concrescence coincide. Low eye swelling occurs at the anterior cardinal angle and is
connected to the anterior marginal ridge.
REMARKS. Oertli placed this species only tentatively in Looneyella, but com-
parison with the holotype of the type species convinces me that L. subtilis is con-
generic. Dr. Oertli kindly sent me specimens of his material for comparison and
enabled me to examine internal characters not available in my material.
Ecologically this is an interesting genus with the Cretaceous species recorded from
sediments considered to be non-marine whilst L. subtilis occurs in truly marine
conditions. Possibly the Cretaceous species lived under brackish-water conditions.
The material from the Upper Bathonian Bradford Clay of Bradford consists of
a single female carapace in the Winwood Collection (10.3659), and two male carapaces
and a juvenile carapace (10.3956-8) from the Fuller's Earth of Bath in the Blake
Collection.
DIMENSIONS. 10.3659, female carapace, length 0-37 mm. ; height 0-22 mm. ;
width 0-21 mm. 10.3957, male carapace, length 0-35 mm.; height o-i8mm.;
width 0-17 mm.
Genus ORTHONOTACYTHERE Alexander 1933
Orthonotacythere sp.
(PL 15, ng. i)
REMARKS. The single specimen is a small carapace having a high caudal process,
a long, straight dorsal margin and a ventral surface triangular in outline. The
lateral surface is weakly reticulate but prominently ornamented by three large
tubercles. The first occurs at the anterior cardinal angle and is an eye tubercle.
The second occurs at the postero-dorsal part of the carapace and is somewhat elongate
in outline, whilst the third is situated directly below the second at the postero- ventral
corner of the carapace. This last tubercle, positioned at the end of a somewhat
thickened ventro-lateral margin, extends slightly beyond the carapace posteriorly.
A weak lateral ridge is present at about valve centre, but does not estend as far as the
terminal margins.
Orthonotacythere nodosa Plumhoff (1963), which appears in the Lias of Germany,
differs from the present species in possessing a much larger number of tubercles.
?0. pulchella Apostolescu (1959) from the Lias of France does not belong to this
genus. Orthonotacythere sp. also differs from the small number of species described
from the Upper Jurassic of the U.S.S.R. (Ljubimova 1955), Germany (Triebel 1941)
and France (Donze 1960).
DIMENSIONS. 10.3993, carapace, length 0-30 mm. ; height 0-15 mm. ; width
0-14 mm.
ENGLISH BATHONIAN OSTRACODA 433
Genus PARARISCUS Oertli 1959
Parariscus bathonicus Oertli
(PI. 15, figs. 2, 8)
1959 Parariscus bathonicus Oertli : 118, pi. 2, figs. 20-29.
REMARKS. Eight complete carapaces of this interesting species are present in the
Blake Collection of Fuller's Earth Ostracoda. Dr. H. J. Oertli kindly sent me
comparative material.
DIMENSIONS. 10.3959, carapace, length 0-43 mm. ; height 0-21 mm. ; width
0-20 mm.
Genus PARACYTHERIDEA Miiller 1894
Paracytheridea? blakei sp. nov.
(PI. 15, figs. 4, 5, 9, 10)
DIAGNOSIS. Carapace small, reticulate, with dorso-median constriction.
Postero- ventral angle swollen, extended backwards beyond margin. Eye node
developed at anterior cardinal angle. Dorsal margin long, straight. Anterior end
high, rounded. Posterior end narrow, triangular, situated high on carapace.
HOLOTYPE. 10.4001, carapace, Fuller's Earth, Bath.
PARATYPES. 10.4014-6, three carapaces, Fuller's Earth, Bath.
DESCRIPTION. Carapace small with coarse reticulate ornamentation. Dorso-
median part of each valve with broad depression. Ventro-lateral margin swollen,
particularly towards the posterior end where, at the postero-ventral angle, there is
a backward projection of the carapace beyond the steeply angled postero-ventral
slope. Dorsal margin long and straight with sharply angled cardinal angles. An
eye node is situated behind the anterior cardinal angle. Anterior end high, rounded.
Posterior end situated high on the carapace, sharply triangular with short, concave
postero-dorsal slope and long, convex postero-ventral slope. Line of greatest length
slightly above the mid-point ; greatest height at the anterior cardinal angle. Greatest
width in the posterior third. Left valve slightly larger than the right, which it
overlaps along the ventral margin. Internal features not seen apart from the muscle
scars which are visible from the exterior : these consist of a slightly crescentic row of
four adductor scars, an antero- ventral, rounded, manibular scar, and an antero-dorsal
antennal scar which is composed of two scars, the first crescentic in shape, the second
rounded and fitting inside the crescent of the first. These muscle scars are situated
low down on the carapace, below the dorso-median depression.
DIMENSIONS. 10.4001, holotype, carapace, length 0-30 mm. ; height 0-16 mm. ;
width 0-14 mm.
REMARKS. Paracytheridea? blakei was named after J. F. Blake, in whose collection
the species was found. A new genus might be preferred, but for the present it is
sufficient to place the species in Paracytheridea on the basis of the external shape.
There is no strong nodose ornamentation as in Paracytheridea? caytonensis Bate
(1965).
434 REVISION OF SOME
Genus CYTHERURA Sars 1866
Cytherura bathonica sp. nov.
(PL 16, figs. 5-8)
DIAGNOSIS. Cytherura with small, elongate carapace, ornamented with slender,
almost parallel, striae. Short, vertical connecting bars between striae present a
slightly reticulate appearance. Posterior extremity situated above the line of the
longitudinal axis.
HOLOTYPE. 10.4017, carapace, Fuller's Earth, Bath.
PARATYPES. 10.4018-9 and 10.4120, eight carapaces. Fuller's Earth, Bath.
DESCRIPTION. Carapace small, elongate, with line of greatest length very slightly
above the mid-point. Greatest height at the extreme anterior end, greatest width
in the posterior third. Shell surface ornamented by slender, almost parallel striae
with connecting vertical bars between. A slightly reticulate appearance is thereby
presented although this is subordinate to the sub-parallel striae. Dorsal and ventral
margins long, straight and almost parallel. Anterior end broadly rounded, posterior
end with narrow triangular margin terminally truncated and set high on the carapace :
the concave postero-ventral slope is much longer than the concave postero-dorsal
slope. Left valve only slightly larger than the right, overlapping it ventrally just
anterior of median and at the cardinal angles. The right valve overlaps the left
along the dorsal margin. Internal features not seen.
DIMENSIONS. 10.4017, carapace, length 0-30 mm. ; height 0-13 mm. ; width
o-io mm. 10.4018, carapace, length 0-31 mm. ; height 0-14 mm. ; width o-n mm.
REMARKS. Cytherura bathonica is a true Cytherura as far as the external details
are concerned. The presence of this species and C. mediojurassica sp. nov. in the
Bathonian indicates that the genus was established by or during the Middle Jurassic.
C. bathonica differs from previously described species in ornamentation although it
comes close to the Recent Cytherura striata Sars (1866) from which it may be distin-
guished by the position of the posterior extremity, which is situated below the
longitudinal axis in C. striata and above in C. bathonica.
Cytherura mediojurassica sp. nov.
(PL 15, fig. 6; PL 16, figs, i, 2)
DIAGNOSIS. Cytherura with reticulate ornamentation. Carapace elongate, dorsal
and ventral margins long, almost parallel but converging slightly towards posterior
end. Short caudal process set high up on carapace.
HOLOTYPE. 10.4121, carapace, Fuller's Earth, Bath.
PARATYPES. 10.4122-38, sixteen carapaces, Fuller's Earth Clay, Bath.
DESCRIPTION. Carapace elongate with long, sub-parallel dorsal and ventral
margins which converge slightly towards the posterior end. Greatest length of
ENGLISH BATHONIAN OSTRACODA 435
carapace through the mid-point whilst the greatest height is at the extreme anterior
end, through the anterior cardinal angle. Greatest width in the posterior third.
Carapace slender in dorsal view. Anterior end rounded, posterior end with short
caudal process set high on the carapace, slightly above the line of greatest length.
Postero-dorsal slope short, concave ; postero- ventral slope long and concave, although
in some specimens it is slightly convex. Termination of caudal process truncated.
Shell surface ornamented by a rather coarse reticulation. Occasionally a weak
longitudinal stria is present in the posterior half of the carapace and extends along
the longitudinal axis as far as the valve centre. This striation is really an exag-
gerated development of the reticulation and in the holotype is present only on the
right valve. Left valve slightly larger than the right in that it overlaps the right
along the ventral margin, especially along its anterior half. Dorsally, however, the
right valve overreaches the left. Internal features not known.
DIMENSIONS. 10.4121, carapace, length 0-33 mm. ; height 0-15 mm. ; width
0-13 mm. 10.4122, carapace, length 0-31 mm. ; height 0-15 mm. ; width 0-12 mm.
REMARKS. Cytherura mediojurassica sp. nov. is easily distinguished from C.
bathonica by its strong reticulate ornamentation, and from Cytherura? liesbergensis
Oertli (1959) by its more elongate carapace in which the dorsal and ventral margins
are sub-parallel. In addition the ornamentation of C.? liesbergensis is weakly
reticulate, with four lateral striae extending along most of the length of the carapace.
These striae are not represented in the present species. Dr. H. J. Oertli kindly sent
me comparative material.
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ENGLISH BATHONIAN OSTRACODA 437
OERTLI, H. J. 19590. Malm-Ostrakoden aus dem schweizerischen Juragebirge. Denkschr.
schweiz. naturf. Ges., Zurich, 83 : 1-44, 7 pis.
1960. Procytheridea parva, nouveau nom. pour Procytheridea minuta Oertli 1959. Rev.
de Micropaleont., Paris, 3 : 70.
1963. Faunes d'ostracodes du Mesozoiqite de France, 57 pp., 90 pis. Leiden.
PECK, R. E. 1951. A new ostracode genus from the Cretaceous Bear River Formation.
/. Paleont., Tulsa, 25 : 575-577, pi. 80.
PETERSON, J. A. 1954. Jurassic Ostracoda from the " Lower Sundance " and Rierdon
Formations, Western Interior United States. /. Paleont., Tulsa, 28 : 153-176, pis. 17-19.
PLUMHOFF, F. 1963. Die Ostracoden des Oberaalenium und tiefen Unterbajocium (Jura)
des Gifhorner Troges, Nordwestdeutschland. Abh. Senckenb., Frankfurt a.M., 503 : i-ioo,
pis. 1-12.
RICHTER, R. 1855. Aus dem thiiringischen Zechstein. Z. dt. geol. Ges. Berlin 7 : 526-533.
1867. Aus dem thiiringischen Zechstein. Z. dt. geol. Ges., Berlin, 19 : 216-236.
SARS, G. O. 1866. Oversigt af Norges marine ostracoder. Vidensk.-Selsk. i Christiania, Fork.,
Oslo, 7 : 1-130.
STEGHAUS, H. 1951. Ostracoden als Leitfossilien im Kimmeridge der Olf elder Wietze und
Fuhrberg bei Hannover. Palaont. Z., Stuttgart, 24 : 201-224, pis. 14-15.
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of Louisiana and Arkansas. /. Paleont., Tulsa, 20 : 362-373, pis. 52, 53.
SYLVESTER-BRADLEY, P. C. 1948. Bathonian ostracods from the Boueti Bed of Langton
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not. Hist. (Geol.), London, 3 : 1-21, pis. 1-4.
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pis. 1-15.
1951. Einige stratigraphisch wertvolle Ostracoden aus dem hoheren Dogger Deutschlands.
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R. H. Bate, B.Sc., Ph.D.,
Department of Palaeontology
BRITISH MUSEUM (NATURAL HISTORY)
LONDON. S.W.7
EXPLANATION OF PLATES
All the specimens illustrated are in the Department of Palaeontology, British
Museum (Natural History). With the exception of PL 5, figs. 5, 6, and PL 16, fig 4,
the photographs were taken by the author with the Stereoscan scanning electron
microscope.
PLATE I
FIG. i. Cytherella symtnetrica Jones. External view, male left valve and lectotype of
Cytherella subovata Jones. IN. 43496. x 75.
FIG. 2. Cytherella symmetrica Jones. External view of lectotype, female left valve.
IN.43503. x 85.
FIG. 3. Cytherelloidea jugosa (Jones). External view of lectotype, female right valve.
IN-43497. x 120.
FIG. 4. Cytherelloidea jugosa (Jones). External view of paralectotype, left side of female
carapace. 1.2311. x 115.
FIG. 5. Bairdia Hilda Jones. Right valve of complete carapace, paralectotype, 10.3608.
x 78.
FIG. 6. Bairdia Hilda Jones. Left valve, lectotype, IN. 4 1951. x 73.
FIG. 7. Bairdia juddiana Jones. Right valve of complete carapace, holotype, IN. 43506.
X 65.
FIG. 8. Lophocythere acutiplicata (Jones & Sherborn). Male right valve, paralectotype,
10.3643. x 95-
Butt. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE i
GEOL. 17, 8
27
PLATE 2
FIG. i. Bairdia jurassica Jones. Left valve of complete carapace, lectotype, 1^43494.
X 73-
FIG. 2. Bairdia jurassica Jones. Right valve of complete carapace, paralectotype.
Specimen figured and described by Jones as B. jurassica var tenuis. 1^43495. x 73.
FIG. 3. Pontocyprella harrisiana (Jones). Left valve of specimen figured and described
by Jones as Macrocypris bradiana. IN. 43500. x 70.
FIG. 4. Schuleridea (Eoschuleridea) subperjorata (Jones). External view, right valve,
lectotype, originally figured as Cytheridea subperforata (in part) by Jones. IN. 43504. x 97.
FIG. 5. Schuleridea (Schuleridea) jonesiana (Bosquet). External view, left valve of
complete carapace originally figured as Cytheridea subperforata (in part) by Jones. IN-4349O.
X 73-
FIG. 6. Praeschuleridea schwageriana (Jones). Right valve of complete carapace,
holotype. ^.43499. Originally figured as Cythere schwageriana Jones, x 99.
FIG. 7. Schuleridea (Eoschuleridea) trigonalis (Jones). Right valve, complete carapace,
holotype. IN. 42373. Originally described as Bairdia trigonalis Jones, x 85.
FIG. 8. Praeschuleridea sp. Right valve of complete female carapace, 10.3619. x 98.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 2
PLATE 3
FIG. i. Glyptocythere guembeliana (Jones). External view, male right valve, lectotype,
IN. 43493. Originally described as Cythere guembeliana by Jones, x 77.
FIG. 2. Glyptocythere guembeliana (Jones). External view, juvenile right valve,
1.1840. Originally described as Cytheridea trapezoidalis Terquem by Jones & Sherborn. x 113.
FIG. 3. Lophocy there bradiana (Jones). Left valve of complete male carapace, lectotype,
IN. 42372. Originally described as Cythere bradiana by Jones, x 101.
FIG. 5. Lophocy there bradiana (Jones). Right valve of complete male carapace, para-
lectotype, 10.3627. x 95.
FIG. 6. Lophocy there bradiana (Jones). Left valve of complete female carapace, para-
lectotype, 10.3628. x 97.
FIGS. 4, 7, 8. Fastigatocythere juglandica (Jones). Fig. 4, external view right valve,
lectotype, IN. 41947, x 75. Figs. 7, 8. Normal pore canals of lectotype, fig. 7 x 2,000,
fig. 8 x 5,000. Originally described as Cythere juglandica Jones.
Butt. Br. Mus. not. Hist. (Geol.) 17, 8.
PLATE 3
PLATE 4
FIG. i. Glyptocythere guembeliana (Jones). Female right valve and lectotype of
Cytheridea pulvinar Jones & Sherborn. 1.1858. x 85.
FIG. 2. Lophocythere bradiana (Jones). Right valve of complete male carapace, 1.1854.
Lectotype of Cytheridea craticula Jones & Sherborn. x 100.
FIG. 3. Lophocythere bradiana (Jones). Male left valve, paralectotype of C. craticula
Jones & Sherborn. 1.1867. x 100.
FIG. 4. Lophocythere bradiana (Jones). Ventral view, female carapace, paralectotype,
10.3629 x 100.
FIG. 5. Bairdia hilda Jones. Internal view, right valve of paralectotype of Bairdia
fullonica Jones & Sherborn. 10.3554. x ?8-
FIG. 6. Bairdia hilda Jones. External view, right valve. Lectotype of B. fullonica
Jones & Sherborn. 1.1873. x 78.
FIG. 7. Micropneumatocythere subconcentrica (Jones). Left valve of lectotype
IN. 43505. Originally described as Cythere subconcentrica Jones, x 100.
FIG. 8. Terquemula blakeana (Jones). Right valve originally described as Cytheridea
ransversiplicata Jones & Sherborn. 1.1828. x 100.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 4
PLATE 5
FIG. i. Terquemula blakeana (Jones). Female left valve, lectotype, IN. 41950. Origi-
nally described as Cythere blakeana Jones, x 100.
FIG. 2. Glyptocythere oscillum (Jones & Sherborn). Male left valve, lectotype of
Cytheridea striblita Jones & Sherborn. 10.3943. x 95.
FIGS. 3, 7. Cythereis cf. blanda Kaye. External view, left valve, IN. 43502 and right
valve of complete carapace, IN. 43491. Both specimens originally described as Cythere
(Cythereis) quadrilatera (Roemer). x 100.
FIGS. 4-6. Metacytheropteron drupaceum (Jones). Left valve from the Blake Collec-
tion, 10.3977. x 100, and left and right views of complete carapace, holotype, IN. 43498.
X 85. Originally described as Cythere drupacea Jones.
FIG. 8. Cythere ? tenella Jones. Right valve of complete carapace, holotype, IN. 43492.
X 100.
FIG. 9. Cytherella fullonica Jones & Sherborn. Left valve of paralectotype, 10.4004.
x 105.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 5
PLATE 6
FIG. i. Cytherella fullonica Jones & Sherborn. Left valve, lectotype, 1.1857, x IO1-
FIGS. 2, 3. Cytherelloidea catenulata (Jones & Sherborn). Right valve, paralectotype,
1.1846, x 100, and left valve, lectotype, 1.1876, x 100. Species originally described as
Cytherella catenulata Jones & Sherborn.
FIGS. 4-7. Cytherelloidea refecta (Jones & Sherborn). Fig. 4, female left valve, 10.3931
paralectotype, x 100. Figs. 5, 6, male left valve, paralectotype, 10.3932, fig. 5, x 98, fig. 6,
anterior ornamentation of same specimen x 5,000. Fig. 7, female right valve, lectotype,
1.1850, x 95.
FIG. 8. Bairdia sherborni sp. nov. Left valve, complete carapace, holotype 10.3913.
X 107.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 6
PLATE 7
FIG. i. Bairdia sherborni sp. nov. Right valve, complete carapace, paratype, 10.3557.
X 96.
FIG. 2. Paracypris terraefullonicae (Jones & Sherborn). Right valve of lectotype of
Macrocypris horatiana Jones & Sherborn. 1.1874. x I0°-
FIG. 3. Monoceratina visceralis (Jones & Sherborn). Left valve, holotype, 1.1830,
X 100. Originally described as Cytheridea visceralis Jones & Sherborn.
FIG. 4. Paracypris terraefullonicae (Jones & Sherborn). Left valve, lectotype, 1.1875.
Originally described as Macrocypris terrae-fullonicae Jones & Sherborn. x 100.
FIG. 5. Monoceratina vulsa (Jones & Sherborn). Damaged left valve, lectotype, 1.1842,
X 100. Originally described as Cytheridea vulsa Jones & Sherborn.
FIG. 6. Oligocythereis fullonica (Jones & Sherborn). Right valve of complete carapace,
lectotype, 1.1871. Originally described as Cythereis fullonica Jones & Sherborn. x in.
FIG. 7. Schuleridea (Eoschuleridea) horatiana (Jones & Sherborn). Male right valve,
lectotype, 1.1852. Originally described as Cytheridea horatiana Jones & Sherborn. x 56.
FIG. 8. Asciocy there obovata (Jones & Sherborn). Right valve of complete carapace,
paralectotype, 10.3938, x 100. Originally described as Cytheridea obovata Jones & Sherborn.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 7
PLATE 8
FIG. i. Eocytheridea sp. Right valve, 10.3906, x 85.
FIGS. 2, 7. Asciocythere obovata (Jones & Sherborn). Left valve of complete carapace,
10.3939, x 100 and part of shell surface showing normal pore canals, x 650.
FIG. 3. Asciocythere obovata (Jones & Sherborn). Right valve, lectotype, 1.1836.
X 100.
FIGS. 4-6. Praeschuleridea subtrigona (Jones & Sherborn). Male right valve, lectotype,
1.1838, x 98; right valve, female carapace, paralectotype, 10.3937, x 100 and left valve, male
carapace, paralectotype, 10.3936. x 100.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 8
PLATE 9
FIGS. 1-8. Hadrocytheridea dolabra (Jones & Sherborn). Figs, i, 4. External and
internal views, female right valve, lectotype, 1.1851, x 100. (Lectotype of Cytheridea dolabra
Jones & Sherborn.) Fig. 2. Female left valve Cytheridea pentagonalis of Jones & Sherborn),
10.3929, x 100. Fig. 3. Internal view male right valve (Cytheridea parallela of Jones &
Sherborn), paralectotype, 10.3926, x 100. Fig. 5. Male left valve, paralectotype, 1.1844
(lectotype of Cytheridea puteolata Jones & Sherborn), x 100. Fig. 6. Female left valve,
paralectotype, 1. 1866 (holotype of Cytheridea pentagonalis Jones & Sherborn), x 100. Fig. 7.
Male right valve, complete carapace, paralectotype, 1.1859 (lectotype of Cytheridea parallela
Jones & Sherborn), x 100. Fig. 8. Right valve, possibly juvenile instar of H. dolabra (lectotype
of Cytheridea ignobilis Jones & Sherborn), 1. 1868, x 100.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
GEOL. 17, 8.
PLATE 10
FIG. i. Hadrocytheridea dolabra (Jones & Sherborn), Dorsal view, male carapace,
paralectotype, 1.1859 (lectotype of Cytheridea parallela Jones & Sherborn). x 100.
FIGS. 2, 7, 8. Caytonidea terraefullonicae (Jones & Sherborn). Dorsal view, female
carapace, paralectotype, 10.3920, x 100; female left valve, lectotype, 1. 1869, x 100 and female
right valve, paralectotype, 10.3919, x 100. Species originally described as Cytheridea terrae-
fullonicae Jones & Sherborn.
FIGS. 3-6. Rectocy there sugillata (Jones & Sherborn). Figs. 3, 6, external and internal
view, left valve lectotype, 1.1855, X 100. Figs. 4, 5, external and internal view, right valve
paralectotype, 10.3930, x 100. Species originally described as Cytheridea sugillata Jones &
Sherborn.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 10
PLATE ii
FIGS. 1-3. Caytonidea terraefullonicae (Jones & Sherborn). Internal view, female left
valve, lectotype, 1. 1869, x 100; internal view x 100 and surface ornamentation x 750 of
female right valve, paralectotype, 10.3919.
FIGS. 4-6. Acanthocythere sphaerulata (Jones & Sherborn). Right valve, complete
female carapace x 100 and enlargement of surface ornamentation x 2,800 and x 500. Holo-
type, 1.1835. Species originally described as Cythere sphaerulata Jones & Sherborn.
Bull. Br. Mus. not. Hist. (Geol.) 17, 8.
PLATE ii
PLATE 12
FIG. i. Glyptocythere persica (Jones & Sherborn) . Right valve, holotype, 1.1834, X IO°-
Originally described as Cytheridea persica Jones & Sherborn.
FIG. 2. Glyptocythere oscillum (Jones & Sherborn). Female right valve, lectotype,
1.1849 (holotype of Cy there oscillum Jones & Sherborn), x 105.
FIG. 3. Fastigatocy there juglandica (Jones & Sherborn). Female left valve, 1.1872,
X 92. Originally described as Cythere juglandica var major by Jones & Sherborn.
FIGS. 4, 6. Lophocy there acutiplicata (Jones & Sherborn). Left valve, female carapace,
lectotype, 1.1863, X 112 and internal view, female right valve, paralectotype, 1.1847, x 103.
FIG. 5. Lophocythere ostreata (Jones & Sherborn). Right valve, holotype, 1.1833, x 85.
Originally described as Cytheridea ostreata Jones & Sherborn.
FIG. 7. Lophocythere fulgurata (Jones & Sherborn). Right valve, lectotype, 1.1832,
X 85. Originally described as Cytheridea fulgurata Jones & Sherborn.
FIG. 8. Lophocythere septicostata Bate. Female right valve, 1.1843, x 100. Originally
described as Cytheridea bradiana Jones & Sherborn.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 12
PLATE 13
FIG. i. Micropneutnatocy there limaciformis (Jones & Sherborn). Right valve,
holotype, 1.1831. Originally described as Cytheridea limaciformis Jones & Sherborn. x 85.
FIG. 2. Progonocy there stilla Sylvester-Bradley. External view, left valve, 10.3657.
X 114.
FIG. 3. Cytheridea aequabilis Jones & Sherborn. Left valve of complete carapace,
holotype, 1.1864. x 80.
FIG. 4. Cythere corrosa Jones & Sherborn. Right valve of complete carapace, holotype,
1.1865. X I0°-
FIG. 5. Cytheridea coarctata Jones & Sherborn. Left valve, holotype, 1.1841. x 100.
FIG. 6. Cytheridea eminula Jones & Sherborn. Left valve, lectotype, 1.1839. x 100.
FIG. 7. Cytheridea spinifastigiata Jones & Sherborn. Left valve, holotype, I.i86i.
X 100.
FIG. 8. Cytheridea punctiputeolata Jones & Sherborn. Right valve, holotype, 1.1856.
x 85.
FIG. 9. Cytheridea spinigyrata Jones & Sherborn. Right valve, holotype, 1. 1860.
X 100.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 13
PLATE 14
FIGS. 1,3. Ektyphocyt here parva Oertli. Left valve, male carapace, 10.3983, x 115 and
right valve, female carapace, 10.3982, x 114.
FIGS. 2, 4, 9. Hekistocy there venosa sp. nov. Left valve, paratype, 10.3996, x 195;
right valve of complete carapace, holotype, 10.3995, x 190, and internal view, left valve,
paratype, 10.4012, x 150.
FIG. 5. Platycythere sp. Right valve of complete carapace, 10.3992, x no.
FIG. 6. Looneyella monticula (Jones). Left valve of complete carapace, holotype,
1.5872, from the Cretaceous Bear River Formation, Wyoming, x 97.
FIGS. 7, 8. Looneyella subtilis Oertli. Left valve of complete male carapace, 10.3957,
x 144, and right valve of complete male carapace, 10.3956, x 144.
Bull. BY. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 14
PLATE 15
FIG. i. Or thonotacy there sp. nov. Right valve of complete carapace, 10.3993, x 144.
FIGS. 2, 8. Parariscus bathonicus Oertli. Left valve of complete carapace, 10.3959,
X 144 and dorsal view of complete carapace, 10.3860, x 144.
FIG. 3. Cytheridea retorrida Jones & Sherborn. Left valve of complete carapace,
10.3923. x 108.
FIGS. 4, 5, 9, 10. Paracytheridea ? blakei sp. nov. Fig. 4, muscle scars of paratype,
10.4015, as viewed externally, x 250; fig. 5, dorsal view, complete carapace, paratype, 10.4016,
X 200; fig. 9, right valve, complete carapace, holotype, 10.4001, x 200 and fig. 10, left valve,
complete carapace, paratype, 10.4014, x 200.
FIG. 6. Cytherura mediojurassica sp. nov. Dorsal view of complete carapace, x 200.
FIG. 7. Hekistocy there venosa sp. nov. Dorsal view of complete carapace, holotype,
10.3995- X 190.
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 15
PLATE 16
FIGS, i, 2. Cytherura tnediojurassica sp. nov. Right valve of complete carapace,
holotype^ 10.4121, x 190 and left valve of complete carapace, paratype, 10.4122, x 195.
FIG. 3. Trachycythere sp. Left valve, 10.3966, x 114.
FIGS. 4, 9. Polycope fungosa sp. nov. Left valve, juvenile carapace, paratype, 10.3953,
X 112, and left valve of complete adult carapace, holotype, 10.3951, x 112.
FIGS. 5-8. Cytherura bathonica sp. nov. Fig. 5, left valve of complete carapace, para-
type, 10.4018, x 195; fig. 8, ornamentation of same specimen, x 1,200; fig. 6, dorsal view of
complete carapace, paratype, 10.4019, x 194; fig. 7, right valve of complete carapace, holotype,
10.4017, x 195-
Bull. Br. Mus. nat. Hist. (Geol.) 17, 8.
PLATE 16
INDEX TO VOLUME XVII
New taxonomic names and the page numbers of the principal references are printed in Bold type. An
asterisk (*) indicates a figure.
Acanthinocyathidae 306, 372
Acanthinocyathus 306, 308
apertus 306, 307*-308; PL 2, figs. 1-4
transiens 306
Acanthistius 87
Acanthocyathus 306
apertus 306, 307
Acanthocythere 411
sphaerulata 411; PL u, figs. 4-6
Acanthuroidei 87, 92
Achomosphaera 174
ramulifera 174-175; PL 10, fig. 7
reticulata 175
sagena 175
Acritarcha 107
Acrogaster 73, 75
brevicostatus 73
daviesi 73
heckeli 73*
parvus 73
Acrognathus 82, 83, 98
Acroporidae 13
Acrostichopteris ruffordi 238
Adnatosphaeridiaceae 171
Adnatosphaeridium 171, 172
chonetum 171-172; PL 10, figs, n, 12
Agastrocyathus 342
Aipichthyidae 62, 92, 97, 100
Aipichthys 49, 62-63, 65, 66, 84, 85, 87, 88, 93, 94,
95. 97. 98, 99
minor 63, 64*, 65*
nuchalis 63
pretiosus 62
velifer 63, 66, 99
Ajacicyathacea 372
Ajacicyathida 305
Ajacicyathidae 308
Ajacicyathus 310, 345, 369
walliseri 312, 314
(Ajacicyathellus) 310
Alataucyathinae 338, 340
Alataucyathus 340, 369, 372
Alataucyathidae 338
(Urcyathus) 334
jaroschevitshi 340
(Alataucyathus) 338, 340
(Anaptyctocyathus) 299, 372
cribripora 340-341, 342, 372; PL n, figs, i, 3
excellenthis 340
flabellus 299, 340, 341-342, 372; PL 10, fig. i
minimiporus 340
verschkhovskajae 340
Albula 54
Alectis 98
Alepocephalus 84
Alethopteris goepperti 228
Algites 221
catenelloides 223
valdensis 221
Allothrissops 84, 85, 91
Alphacyathus 305, 310, 369
annularis 305
cf. annularis 305-306; PL i, fig. 2
Alveolites heteropora 12
Amphiblestrella 20
elegans 20-21, 41; PL 4, figs. 1-3
Amphiblestrum 21
elegans 20
Anabantidae 95
Anabantoidei 89, 93
Anasca 13
Anemia 232
adiantifolia 232
cuneata 232
Anomalopidae 68, 95, 96
Anomozamites schaumburgense 242
Anoplogaster 68
Anoplogasteridae 68, 99
Antigonia 94, 96
Aptea polymorpha 171
cf. polymorpha 170, 171
Apteodinium 130
granulatum 121, 130-131; PL 3, figs. 5, 6
Archaeocyatha 342, 359
Archaeocyathellus 308, 310
(Stapicyathus) 308
stapipora 308, 309, 369; PL i, fig. 5
yukonensis 308
Archaeocyathidae 346
Archaeocyathus 344, 346, 372
440
INDEX
acutus 312
atlanticus 345*, 346
dissepimentalis 358, 359
floreus 310
ichnusae 318
infundibulum 318
irregularis 348
magnipora 311
retesepta 364, 366
retevallum 314, 315
retezona 316
robustus 311
stapipora 308, 309
subacutus 312
trachealis 318
tracheodentatus 318, 319
tubavallum 318
(Protocyathus) stapipora 309
Archaeolynthus 302
porosus 303
Archaeopharetra 312, 342
Ardrossacyathus 354
Areoligeraceae 166
Argentina 54, 86
Arripidae 87
Arripis 87
Asciocythere 404
acuminata 404
lacunosa 404
o&owato 404-405*; PI. 7, fig. 8; PI 8, figs. 2, 3, 7
Ascodinium hialinium no
Ascophora 13
Aspidistes 235
sewardi 235, 236-237*; PI. 6, figs. 3, 4
thomasi 235, 236
Aspidium 235
Asterocyathus 325
Asterotumulus 338
Aulacephalodon 286
Aulolepididae 51, 83
Aulolepis 53-54, 55, 63, 83, 84, 86, 87, 93, 98, 99
typus 52*, 53*
Aulopodidae 81
Aulopus 54, 68, 81, 83, 84, 86, 88, 93, 98
Bactrellaria 24
rugica 24-25, 41; PI. 2, figs. 9-14
Bairdia 383, 397
fullonica 383, 397; PI. 4, figs. 5, 6
harrisiana 385
AiWa 383, 384, 385, 397-398; PI. i, figs. 5, 6;
PI. 4, figs. 5, 6
juddiana 383-384, 395; PI. i, fig. 7
jurassica 384-385; PI. 2, figs, i, 2
tennis 384, 385; PI. 2, fig. 2
sherborni 398; PI. 6, fig. 8; PI. 7, fig. i
trigonalis 386, 395; PI. 2, fig. 7
Bairdiacea 383, 397
Biardiidae 383, 397
Baltisphaeridium armatum 153
bifidum 165
/ercuf 159
longifurcatum 158
multifurcatum 152
pseudohystrichodinium 163
Barroisina trifossata 37, 38
Barysoma 290
Bathy master 91
Bathysoma 49, 78-81, 96, 97, 100
lutkeni 78, 79*, 81, 97
Beaufort series 291
Becklesia 244
anomala 244-245*, 248
sulcata 247*-248, 249; PI. 2, fig. 6
Bedfordcyathus 299, 355, 359
irregularis 359
Beisselina 39, 40
flabellata 40
5/nato 39-40, 41; PI. 8, figs. 3, 4
Beisselinopsis 40
flabellata 40, 41; PI. 8, figs, i, 2
marginata 40
Belontiidae 95
Bennettitales 211
Berenicea 15
sp. 15, 41
Berycidae 68, 88, 94, 95, 96, 99
Beryciformes 49, 56, 81, 83, 86, 87, 88, 92, 95, 96,
97. 98, 99
Berycoidei 49, 63, 67-69, 82, 84, 86, 87, 92, 93, 94,
95. 96, 97. 99, ioo
Berycopsis 57-58, 59, 92
elegans 57*, 58
geramanus 58
Beryx 94
Biflustra 20, 22
meudonensis 26
radula 21
simplex 20
tesselata 25, 26
unipora 19
Brama 87
Bramidae 87
Branchiostegidae 87
Branchiostegus 87
Bronchocyathidae 317-318, 319, 372
Bronchocyathus 317, 318, 319, 372
dentatus 321
trachealis 318
tracheodentatus 319
Brycon 84
Bryophyta 210, 211, 218
? Bryophyte leaf 218, 221, 222*
Bythocypris winwoodiana 395
Bythocytheridae 399
Callaiosphaeridium 151
asymmetricum 152; PI. 6, fig. 6
INDEX
441
Cambrocyathellus 354
Cambrocyathus 354, 356, 370
profundus 357*, 358
Camptocythere lincolnensis 410
Canningia 140
rotundata 141
Cannosphaeropsis 172
choneta 171
Caproberyx 76-77, 78, 94, 95
pharsus 77
polydesmus 77
superbus 76*
Caproidae 93, 96
Capros 96
Carangidae 92
Carpodinium 129
obliquicostatum 129; PI. i, figs. 3, 4
Cassandra 82
Gassiculosphaeridia 141
reticulata 141, 142,; PI. 3, fig. 7; PI. 4, fig. 3
Castanopora 8, 12
guascoi 8
jurassica 7-8, n, 14, 19, 31, 37, 41, PI. 7, figs.
4-7
Catenella 223
opuntia 223
Cave Sandstone 293
Caytonidea 410, 424, 426
faveolata 410, 411
terraefullonicae 410-411; PI. 10, figs. 2, 7, 8;
PI. ii, figs. 1-3
Cellaria 9
smithi 9
Cellepora 9
complanata 29
davaiacensis 9
polythele 9
ricata 30
(Dermatopora) faujasi 8
(Discopora) subgranulata 30
koninckiana 6
Centrarchidae 92
Centroberyx affinis 68
Centropomidae 87, 92
Centropomus 87
Ceratocorys 136
veligera 136
Chaetodon 87
Chaetodontidae 87, 92
Chanares Formation 281
Chanaria 281, 284
Channiformes 89, 93, 95, 96
Chanos 54
Char a knowltoni 214, 215
medicaginula 217
Characeae 217
Charales 214, 217
Charixa vennensis 13
Cheilanthites gopperti 229
Cheilostomata 3, 4, 9, 12, 13, 14
Chelmo 87
Chinle Formation 290
Chlamydophorella 156
wyei 155
Chlorophthalmus 81
Chlorophyceae 214
Chondrus crispus 223
Chouberticyathus 342
Chrysomonadinae ? 140
Chytroeisphaeridia 140
chytroeides 141
euteiches 141; PI. 3, figs. 8, 9
sp. 141
Cichlidae 87
Girconitella 214-215
knowltoni 215-216*-217; PI. i, figs. 1-5
Circulodinium deflandrei 166
Cirrhitidae 87
Cirrhitus 87
Cisternifera 9
Cladocopida 427
Cladocopina 427
Cladograpsus capillaris 185
linearis 185, 186
Cladophlebis albertsii 237
browniana 238
dunkeri 238
longipennis 237, 238
(Klukia) dunkeri 238
Claruscyathus 363
Cleistosphaeridium 152, 157
ancoriferum 155
armatum no, 153-154; PI. 8, figs, i, 2, 12
heteracanthum 152; PI. 7, fig. 8
huguonioti 109, 155-156; PI. 7, fig. 10
var. pertusum 156-157; PI. 7, figs. 6, 7, 9
multifurcatum no, 152-153; PI. 8, figs. 7, 10
polypes 154, 155; PI. 6, figs. 7, 8
var. clavulum 154-155; PI. 6, figs. 9, 10
? aciculare 158; PI. 6, figs, u, 12
? flexuosum 157; PI. 7, fig. 4
? parvum 157-158; PI. 7, figs, n, 12
Climacograptus 192
minimus 191-193; PI. 4, fig. 3
minutus 191*, 192*, 193-194; PI. 4, figs, i, 2
scalaris miserabilis 193 ; PI. 4, fig. 2
tangshanensis linearis 193
spp. 1 88
Clonograptus 204
Clupavidae 86, 98
Clupavus 84, 85
Coilostega 13, 22
Cometodinium obscurum 137
Coniferales 211
Copleicyathus 344
confertus 349
Coronifera 161
oceanica 161, 162; PI. 8, figs. 8, n
442
INDEX
Corynoides calicularis 195
Coscinocyathus 325, 329, 330, 369
annulatus 337
cellularis 330
cribripora 340
equivallum 326
papillatus 334
papillipora 338, 339
peter si 331
quadratus 328
ye/i/er 332
rhyacoensis 329
rugosus 325
textilis 329
unilinearis 330; PI. 13, fig. 4
Coscinopleura 29
lamourouxi 29
sp. 29, 41
Coscinoptycta 325, 330
convoluta 330
Coscionteichus 325
Cribroperidinium 121, 124, 125, 128
edwardsi 127, 128
intricatum 109, no, 125-126*-127*-128;
PI. 2, figs. 1-3
muderongensis 127, 128
orthoceras 121, 127, 128-129, 134*
sepimentum 125
Crisisina 17
carinata 17, 41; PI. i, figs. 4, 5
Cristivomer 84
Cryptograptus 194
tricornis 191*, 194-195; PI. 4, figs. 4, 5, 6
schaeferi 195
Cryptostomata 14
Ctenostomata 14
Ctenothrissa 54-56, 63, 81, 83, 84, 86, 87, 93, 98,
99
microcephala 55*
radians 55
signifer 54*. 55
vexillifer 54
Ctenothrissidae 54, 83
Ctenothrissiformes 49, 50-51, 83, 86, 98, 99
Cyathocricus 299, 318-319, 369, 372
annulispinosus 318, 320, 322, 369
dentatus 318, 319, 321*-322; PI. 6, figs. 1-3
tracheodentatus 318, 319-320*, 322; PI. 5, figs.
4, 5 ; PI. 6, fig. 4
Cycadales 210, 211, 242
C>»fls 246
revoluta 246, 248
Cyclocyathellidae 317
Cyclocyathus 318, 319, 372
Cyclonephelium 166, 168, 171
compactum 167
distinctum 166-167, 169*, 170; PI. n, figs. 6, 7,
8, 10
eisenacki 166, 169*, 170-171; PI. 8, figs. 3, 4;
PI. 9, fig- 4
membraniphorum 166, 167-168; PI. u, fig. 9
paucispinum 166, 170; PI. 9, figs, i, 2
vannophorum 166, 168, 169*, 170; PI. 9, fig. 3
PI. n, figs, n, 12
uttadalicum 168
Cyclostomata 9, 15
Cymatiosphaera cingulata 173
Cynidiognathus longiceps 291
Cynognathus Zone 257, 285, 290, 291, 292, 293
Cypridacea 384, 399
Cyrtograpsus murchisoni 202
Cyrtograptus centtrifugus 202, 203
murchisoni 202-2O3*, PI. 5, figs 8, 9
bohemicus 202
crassiusculus 203
rigidus 203
Cytheracea 385, 399
Cythere 394
blakeana 393
bradiana 390, 391
corrosa 424, 426 ; PI. 13, fig. 4
drupacea 393, 394, 427
guembeliana 388, 412
juglandice 389
major 389, 412
minor 395
monticula 431
oscillum 412, 413; PI. 12, fig. 2
schwageriana 387; PI. 2, fig. 6.
speciosa 395
sphaerulata 411
subconcentrica 392
? teneWo 394; PI. 5, fig. 8
trapezioides 395
walfordiana 395
(Cythereis) quadrilatera 394
Cythereis 394
blanda 394
cf. Wawrfa 394; PI. 5, figs. 3, 7
fullonica 401
walfordiana 395
Cytherella 380, 381, 395
catenulata 396
fullonica 395; PL 5, fig. 9; PI- 6, fig. i
jugosa 381
subovate 380, 381
suprajurassica 381
symmetrica 380-381; PI. i, figs, i, 2
Cytherellidae 380, 395
Cytherelloidea 381, 396
catenulata 396; PI. 6, figs. 2, 3
eastfieldensis 382
;«£osa 381-382*; PI. i, figs. 3, 4
paraweberi 382*
ye/ec/a 396-397; PI. 6, figs 4-7
weberi 382
Cytheridae acutiplicata 416
aequabilis 424; PI. 13, fig. 3
INDEX
443
bicarinata 395, 420, 421
blakeana 393, 423
bradiana 391*, 418
coarctata 424; PL 13, fig. 5
craticula 390, 391*, 392, 418; PL 4, figs. 2, 3
dolabra 406, 407, 408
egregia 393. 395
eminula 425; PL 13, fig. 6
eximia 395
fulgarata 419
horatiana 402
ignobilis 407, 408, 409; PL 9, fig. 8
limaciformis 422
obovata 404
ostreata 420
parallels, 407, 408; PL 9, fig. 3; PL 10, fig. i
pentagonalis 407, 408; PL 9, figs. 2, 6
persica 415
politula 395
pulvinar 388, 412; PL 4, fig. i
punctiputeolata 409, 425; PL 13, fig. 8
£«ra 395
puteolata 407, 408; PL 9, fig. 5
refecta 396
renoides 395
retorrida 394, 425*-426*; PL 15, fig. 3
rugifera 395
sedata 395
spinifastigiata 426; PL 13, fig. 7
spinigyrata 426-427; PL 13, fig. 9
striblita 412, 413; PL 5, fig. 2
subeminula 395
subperforata 385; PL 2, fig. 4
subtrigona 405
sugillata 409
terraefullonicae 410
transversiplicata 393, 423
trapezoidalis 388, 412
ventrosa 395
visceralis 399
vufca 400
winwoodiana 395
Cytherideidae 406
Cytherideinae 406
Cytheropteron jonesanum 393, 394
Cytherura 434
bathonica 434; PL 16, figs. 5-8
? liesbergensis 435
mediojurassica 434-435; PL 15, fig. 6;
PL 1 6, figs, i, 2
striata 434
Cytheruridae 393, 431
Cyttus 96
Dactylopteriformes 96
Daptocephalus 286
Decurtaria 37
allecta 38
cornuta 37, 38
cf. cornuta 37-38, 41; PL 7, figs, i, 2
Deflandrea no
Dendrocyathus 354
Dendrograptus lentus 204
Dermogenys go
Description of sections
Compton Bay, Isle of Wight 112, 113*
Devon 115, 117*
Escalles Borehole, Pas de Calais 115, 118*
Fetcham Mill, Surrey in*, 112
Hunstanton, Norfolk 115, 116*
Saskatchewan 115, 119*
Speeton, Yorkshire 112, 114*
Texas 115
Diademodon 259, 292
polyphagus 292
rhodesiensid 292
Diastopora 9
lamellosa 5
ranvilliana 9
Dicellograpsus elegans 187
moffatensis 189
Dicellograptus 188, 190
anceps 189
duvaricatus 189, 190
elegans 187-188*, 190; PL 3, fig. i
moffatensis 188*, 189-190; PL 3, figs. 5, 6
alabamensis 190
morrisii 188, 190
pumilus 1 88
vagus 190
Dicentrarchus 87
Dichopteris delicatula 236
sp. cf. £>. laevigata 240
Dicranograptus 190, 191
brevicaulis 190
clingani 190-191*; PL 3, figs. 2, 3, 4
Dictyocyathidae 342, 344
Dictyocyathus 342, 344, 370
annularis 305
graphicus 346, 348
Dictyophyllum roemeri 228
Dictyopyxidia 141
Didymograpsus elegans 187
Didymograpsus moffatensis 189
Dimorphostylus 32, 33
tetrastichus 32, 33
Dinanomodon 286
Diniferophycidae 120
Dinodontosaurus 281, 284
turpior 283, 284
Dinophyceae 120
Dinopterygidae 62, 66-67, 92, 98
Dinopterygoidei 62, 86, 87, 88, 97, 99
Dinopteryx 67, 68, 84, 87, 88, 93, 95, 99
spinosus 66
Diphyes 161
Diplograpsus foliaceus 194
minimus 191, 192*, 193
444 INDEX
minutus 192, 193
pristis 192, 193
tricornis 194
Diplograptidae 194
Diplopholeos 27
Dipteridaceae 228
Diretmidae 68, 88, 99
Diretmus 68, 88, 99
Diversograptus ? capillaris 198, 199*. 200
Doidyx 142
Dokidocyathidae 305, 372
Dokidocyathus 305
Drepane 87
Esienackia eras sit abulata 136
Ektyphocy there 430
parva 430; PL 14, figs, i, 3
Ellipsoidictyum 141
cinctum 141
Ellipsodinium 129-130
rugulosum 130, 139*; PL 3, fig. I
Eops 54, 63, 83, 84, 85, 86, 88, 98
Emydops 285
Endothiodon Zone 258
Entalophora g, 16
benedeniana 16, 41
madreporacea var. benedeni 16
Eocytheridea 406
faveolata 406
sp. 406; PL 8, fig. i
Epelidosphaeridia 142
spinosa 143; PL 3, figs. 10-12
Ephippidae 87, 98
Ephippus 87
Equisetales 211, 227
Equisetites 227-228
/ye//i 227
Er-ma-ying Formation 290
Erismacoscinidae 325
Erismacoscinus 325, 332, 369
cancellatus 332
cellularis 330-331; PL 8, figs, i, 4
equivallum 326-327, 328, 331; PL 7, figs. 1-3, 5
marocanus 325
papillatus 334
petersi 331-332, 333; PL 10, figs. 4, 5
quadratus 328-329; PL 7, fig. 4; PL 8, fig. 5
retifer 332-333; PL 10, figs. 2, 3
rugosus 325-326, 328, 331; PL 8, figs. 2, 3
textilis 329-330; PL 9, figs. 1-4
(Pluralicoscinus] 325
Erythrosuchus 291
Escarpment Grit 258, 293
Escharidae 9
Eschar a 9, 24
bellona 35
caecilia 36
cepha 34
gaimardi 24
34
40
ranvilliana 9
striata 39
Escharifora flabellata 40
Escharinella simplex 30
Ethmocoscinus 338-339, 369
papillipora 339; PI. 3, fig. 3
Ethmocyathidae 299, 322-323, 372
Ethmocyathus 322, 323, 369, 372
lineatus 323-324*; PI. 5, figs. 1-3
Ethmophyllidae 317
Ethmophyllum 313, 314, 370
dentatum 318, 319, 321
flexum 315, 316
poletevae 315
vermiculatum 315
Euparkeria 291
Exellia 97, 98, 100
Exochosphaeridium 131, 162, 166
" hirsutum " group 164, 165
phragmites 163; PI. 7, fig. 5
pseudohystrichodinium 163-164; PI. n, figs. 4, 5
spinosum 166
var. deflandrei 165, 166
striolatum 162, 163, 164, 165
var. truncatum no, 164-166; PI. 7, figs. 1-3
(Hystrichosphaeridium) cf. hirsutum 165
Fastigatocythere 389, 412
juglandica 389, 412; PI. 3, figs. 4, 7, 8; PI. 12,
fig. 3
Filicales 228
Flindersicyathidae 344, 372
Flindersicyathus 344-346, 350, 370, 371, 372
(Flindersicyathus) 344, 345, 348, 349, 370, 371,
372
circliporus 346
contractus 346
decipiens 344, 348
graphicus 346-347*-348, 363; PI. 12, figs. 3, 4, 5
irregularis 346, 348-349; PI. 14, fig. 4
latiloculatus 346
macdonnelli 346
mo/or 346, 349-350; PL 14, fig. 2
multifidus 346
yete 346, 348
simplex 346
speciosus 346, 350; PI. 14, figs, i, 3
tabulatus 346
(Pycnoidocyathus) 344, 345, 350-351, 352, 354,
37°. 3?i. 372
maximipora 346, 352-353; PI. 15, fig. 3
parvulus 346
ptychophragma 346
simplex 352; PL 15, fig. 2
synapticulosus 346, 350, 351; PL 15, fig. i
vicinisepta 346, 353-354; PL 15, fig. 4
Floridina 31
INDEX
445
Floridinella 31
scutata 31; PI. 4, fig. 7
" Flustra " flabelliformis 4-5
Flustrella 20
baculina 20
convexa 25, 26
irregularis 21
marginata 25
simplex 19
Flustrellaria incrassata 23
Fromeaceae 140
Fromea 140
amphora 140; PI. 3, figs. 2, 3
Frurionella 38, 39
europaea 39; PI. 8, fig. 8
fertilis 3, n, 38-39, 41; PI. 8, figs. 5-7
Gabrielsocyathus 363
deasensis 364
gabrielsensis 364
poolensis 364
Gasterosteiformes 96
Gaterin 87
Geniculicyathus 325
Gephyroberyx 68
Gesoriacula 431
Gibberichthyidae 68, 96
Gibberichthys 68
Ginkgoales 210, 211, 248
Gleichenites cycadina 238
Glossograptus 194
Glyptocythere 388, 389, 412
costata 416
guembeliana 388-389, 412, 416; PI. 3, figs, i, 2;
PI- 4, fig- i
juglandica 389
oscillum 412-415 [414*]; PI. 5, fig. 2; PI. 12,
fig. 2
persica 415*-416; PI. 12, fig. i
rudimenta 413, 414*, 415
tuberodentina 416
Gnathoberyx 74-75
stigmosus 74
Gonyaulacysta 120, 121, 125, 128, 132
ambigua 124
cf. ambigua 120
apionis 127, 128
cassidata 120
delicata 123*-124; PI. i, figs. 7, 8
diaphinis 127, 128
edwardsi 125
exilicristata no, 121, 122*, 125; PI. i, figs.
i, 2
fetchamensis 120-121
muderongensis 125
obscura 128
orthoceras 125
wetzeli 128
120
Sp. A. 122*, 124-5; PI. i, figs. 9, 10
Gonyaulacystaceae 120
Gonyaulax 125
edwardsi 128
helicoidea subsp. cassidata 120
muderongensis 128
orthoceras 109, 128
sp. no, 127
Grammascosoecia 19
dichotoma 19
Grammicolepidae 96
Graptolithus becki 195
clingani 195
hisingeri 197
intermedius 196
millepeda 195
nilssoni 197
tenuis 197
Gymnospermae 242
Gyrogona 217
medicaginula 217-218
Hadrocytheridea 379, 406
dolabra 407-409 [408*], 425; PI. 9, figs. 1-8;
PI. 10, fig. i
Hagenowinella 23
cf. incrassata 23-24, 41 ; PI. 2, figs. 1-3
Hausmannia 228
dichotoma 228; PI. 3, figs, i, 2
pelletieri 228
Hekistocythere 379, 428, 430
venosa 428-429*^430; PI. 14, figs. 2, 4, 9;
PI- 15, fig- 7
Heliodinium voigti 175
Heniochus 87
Hepaticae 218-219
Hepaticites 219, 220, 221
amauros 223
arcuatus 220
haiburnensis 220
laevis 220
224
rosenkrantzi 223
ruffordi 220, 223; PI. 2, figs. 4, 5
zeilleri 219-220, 223; PL 2, figs. 1-3
Heterocella 27
Heterocrisina 16
abbottii 16
communis 16-17, 41
Heteropora reticulata 18
Hexasphaera asymmetrica 152
Hiodon 84
Hippothoa 9
simplex 13
Histiocysta 138
palla 138-139*-140; PI. i, figs. 5, 6
Histiopterus 87
Holocentridae 67, 68, 75-76, 77, 88, 94, 95, 99
Holocentrus 68, 75, 76
446
ascensionis, 68, 76
Homocytheridea 424
Homonotichthys 60-61, 88, 94, 95, 99
dorsalis 60, 6 1
pulchellns 60*. 6 1
rotundus 61
Hoplopteryx 69-72, 73, 75, 94, 95
antiquus 70
gephyrognathus 70, 71
lewesiensis 70*, 71*, 72
lewisi 71
macracanthus 70, 71
siwws 70, 71
spinulosus 71
syriacus 71, 72
Hoplostethus 68, 70, 72, 74, 75, 94
mediterraneus 67*, 68, 69
Howesia 291
Hymenopteris psilotoides 238
Hynnis 98
Hystrichodiniwn 175
dasys 175-176; PI. 10, figs. 8, 9
pulchrum 109
uoi^rt 109, 175; PI. 10, figs. 6, 10
Hystrichokolpoma 159
/m?# 159-160; PI. 9, figs. 5, 6, 7
Hystrichosphaeraceae 172
Hystrichosphaera no, 172
cingulata var. cingulata 173
var. perforata 173
var. polygonalis 174
var. recticulata 173-174
crassimurata 174
crassipellis 174
furcata 172, 173
ramosa var. gracilis 172
var. multibrevis 173; PI. 10, figs. 3, 4
var. ramosa 172; PI. 10, figs, i, 2, 5
var. reticulata 173
cf. rowosa 174
Hystrichosphaeridiaceae 143
Hystrichosphaeridium 143
ancoriferum 155
anthophorum 147
armatum 153
asterigerum no
asymwe^ncwm 152
bowerbanki 145; PI. 5, fig. 9
complex 146
costatum 146
deanei 144; PI. 4, fig. I
145-146; PI. 4, figs. 2, 6, 7
159
heteracanthum 152
hirsutum 165
huguonioti 155
longifurcatum no, 158
mantelli 109, 145; PI. 4, fig. 9
multifurcatum 152, 153
INDEX
parvispinum 161
polytrichum no
pseudohystrichodinium 163
radiculatum 144-145; PI. 4, fig. 8
readei 144
recurvatum 151, 154
subsp. polypes 154
reniforme 148
cf. salpingophorum 109
siphoniphorum 148
spinosum var. deflandrei 166
stellatum 144
cf. striolatum no
tubiferum 143-144; PI. 5, figs. 5, 8
sp. 22. 1 10
sp. 23. in
I chthyodectes 84
Idmidronea 16
macilenta 16, 41
Idmonea abbottii 16
communis 16
disticha 16
lichenoides 18
macilenta 16
pseudodisticha 16
ramosa 16
(Tubigera) antiqua 16
Irregularia 342, 370
I schigualastia 257, 285, 290
Ischigualasto Formation 290
Kannemeyeria 285, 290, 291, 292
Kannemeyeriidae 290
Kansius sternbergi 78
Kawinga Formation 293
Kingori Sandstone 293
Kinkelinella 430
tenuicostati 430
Kirtonellinae 430
Kistecephalus Zone 258, 285, 286, 293
Korojonia dubiosa 109
Korsogasteridae 68
Kuhliidae 87
Kuhlia 87, 91
Kyphosus 87
Kyphosidae 87, 92
Labiostomella 14
Lampridae 98
Lampridiformes 49, 78, 81, 96, 97, 98, 100
Lampridoidei 78, 97
Lampris 78, 81, 97
Lateolabrax 87
Leckenbya valdensis 238
Lepidorbitoides 10
Leptograptus 185, 187
capillaris 185-186; PI. i, figs, i, 2
flaccidus 186
INDEX
447
arcuatus 186
Leptolepis 84, 91, 98
coryphaenoides 85
dubia 85
normandica 85
Leptosomus 82
Lethrinidae 87
Lethrinus 87
Lingulodinium 165
Lisso&ery* 72, 73, 75, 94, 95
dayi 72*
Litosphaeridium 148
siphoniphorum 148-149*-150; PI. 6, figs. 3, 4
Lobotes 87
Lobotidae 87
Loculicyathus (Loculicyathus) 310, 372
Loculicyathus (Loculicyathellus) 299, 310, 372
/Zoyews 310-311; PI. 3, figs, i, 2, 4
Lonchopteris mantelli 239
Looneyella 431
monticula 431; PI. 14, fig. 6
quadrispina 431
subtilis 431-432; PI. 14, figs, 7, 8
Lophocythere 389-390, 416; 417, 418, 421
acutiplicata 416-417; PI. I, fig. 8; PI. 12, figs.
4,6
bradiana 390-391*-392, 416, 417, 418; PI. 3,
figs, 3, 5, 6; PI. 4, figs. 2, 3, 4
fulgurata 419-420; PI. 12, fig. 7
multicostata 392
ostreata 390, 419, 420-421*; PI. 12, fig. 5
scabra 421
septicostata 391, 418; PI. 12, fig. 8
sp. 418*-419
Lower Cambrian archaeocyathid Stratigraphy
368-371, 372
Australia 368, 369, 370, 371
Morocco 369
Sardinia 369, 370
Siberian platform 368, 369, 370
Southern Siberia 368, 369, 370
Luangwa drysdalli 259, 292
Luciocephalus 89
Lunulites 34
sp. ii, 34, 41
Lutjanidae 87
Lut janus 87
Lycopodiales 224
Lystrosaurus 285
Lystrosaurus Zone 285, 291, 293
Macristiella perlucens 51
Macristiidae 51
Macristium 50
chavesi 50
Macrocypris bradiana 385; PL 2, fig. 3
horatiana 399; PI. 7, fig. 2
terraefullonica 399
Madumabisa Mudstone 293
Malacostega 13, 21, 22
Manda Formation 290, 291, 292, 293
Marchantiales 218, 220
Marchantiolites 219
Marchantites 218, 219, 220
hallei 220
zeilleri 219
Matoniaceae 228
Matonidium 228
goepperti 228
Medialuna 87
Melamphaeidae 68
Melamphaes 68
Membranilarnax cf. pterospermoides 140
Membranipora 14, 19
bipunctata 8
constricta 13; PI. 7, fig. 8
elliptica 13
fragilis 13
genucia 19, 20
jurassica 3, 4, 6, 7
neocomiensis 13
? obliqua 13
simplex 20
unipora 11, 19-20, 41; PI. 2, figs. 7, 8
Membraniporae 13, 20
Mene 80, 81, 97, 98, 100
maculata 80*, 81
oblongus 80
phosphaticus 97
rhombeus 80
Menidae 80, 100
Mesosuchus 291
Metacoscinidae 363, 367
Metacoscinus 363-364, 370
insigne 364
reteseptatus 363, 364-366; PI. 17, figs. 1-4
Metacyathidae 354
Metacyathus 299, 355, 356
irregularis 355, 359
tey/on 355, 360
Metacytheropteron 393, 427
drupaceum 393-394; PI. 5, figs. 4-6
sp- 393
Metafungia 354, 358, 361-362, 370
reticulata 361, 362-363; PI. 18, figs. 1-3
Metaldetes 299, 348, 354, 355-356, 358, 370
columbianus 356
conicus 358, 359
cylindricus 355, 356
dissepimentalis 356, 358-359, 361 ; PI. 16, figs.
i-3
dissutus 356
graphica 346, 348
irregularis 356, 359-360, 361; PI. 16, fig. 4
proteus 356, 358
ramulosus 356
solidus 356
spiralis 358
448 INDEX
superbus 358
taylori 358, 360-361; PI. 13, figs. 1-3
Micrhystridium ambiguum 109
veligerum 136
sp. ? 138, 140
Microdiniaceae 132
Microdinium 132, 135, 136, 137
? crinitum 137; PI. 2, figs. 7, 8
distinctum 133, 134*, 135; PI. 2, figs. 9-11
echinatum 133
irregular e 136
ornatum 132, 133
cf. ornatum 132-133, 134*; PI. 4, fig. 5
setosum 133, 134*; PI. 2, fig. 4
variospinum 134*, 135; PI. 2, figs. 5, 6
veligerum 136-137; PI. 3, fig. 4; PI. 4, fig. 4
Micropneumatocythere 392, 422
limaciformis 422-423; PI. 13, fig. i
quadrata 422, 423
subconcentrica 392; PI. 4, fig. 7
Micropora 32, 33
rw^'ca 32, 33
subgranulata 30
transversa 32-33, 41; PI. 7, figs. 9-12
Microporidae 13, 33
Mohria 232
Molteno Beds 291, 293
Monocentridae 68, 69, 95, 96
Monocentris 63, 68, 69, 87, 88, 89, 99
japonicus 69*
Monoceratina 399, 400
ungulina 400
visceralis 399-400; PI. 7, fig. 3
im/sa 400-401 ; PI. 7, fig. 5
Monocyathida 302
Monocyathidae 302
Monocyathus 302-303, 305, 369
absolutus 302
bilateralis 302
contractus 302
copulatus 302
kuzneskii 302
lebedevae 302
irregularis 304
macrospinosus 302
mellifer 302-304; PI. i, fig. 4
nalivkini 302, 303
operculatus 302
partibus 303
polar is 303
porosus 302, 303; PI. i, fig. 3
robustus 303
sibiricus 303
simplex 303
sparsipora 303
spinosus 303
tenuimurus 303
to#t 303
unimurus 303
Monodactylidae 87, 92
Monodactylus 87
Monograptus 198
attenuatus 198, 200
capillaris 198
clingani 195-196; PI. 5, figs. 1-5
communis 197
gemmatus 198, 199*, 200
intermedius 196*-198; PI. 5, fig. 6
involutus 197
spiralis 203
sp. (atazms or acinaces type) 192
(Rastrites) gemmatus 198
M or one 87
Mugil 88, 92
Multicavea 19
Multicrescis 18, 19
laxata 7, n, 14, 18-19, 31, 41 ; PI. i, figs. 6-8
Musci 218
Myctophidae 83, 99
Myctophoidei 49, 50, 81-83, 86, 88, 98, 99
Myctophum 81
Myodocopida 427
Myosaurus 285
Myripristis 68, 75, 76, 77
adustus 75*
Naiadita lanceolate 218
Nandidae 89, 93
Nathorstia valdensis 238
Nemagrapsus capillaris 185
Nematonotus 63, 68, 82, 83, 84, 85, 86, 88, 93, 98
fcottae 82*, 83
longispinus 82*
Nemipteridae 87
Nemipterus 87
Neocyttus 96
Neomiodon 221
Neoscopelus 81
Neothunnus 89
Nilssonia 242, 244
revoluta 244
schaumburgensis 237*, 242-243*-244
Nitella 215, 217
flexilis 21 7
hyalina 217
polygyra 217
Nitellaceae 217
Nitellites 214
sahnii 217
sp.
Ntawere Formation 257, 258, 259, 262, 283, 284,
286, 290, 292, 293
Okulitchicyathus 354
Oligocythereis 401
fullonica 401-402; PI. 7, fig. 6
Oligosphaeridium 146
anthophorum 147-148; PL 5, figs. 1-3
INDEX
449
complex no, 146-147; PI. 5, figs. 6, 7
perforatum 148
prolixispinosum 147; PI. 5, fig. 4
pulcherrimum 148
reniforme 148; PI. 6, fig. i
reticulatum 147
Omosoma 58-59, 92
intermedium 58
pulchellum 58
sahelalmae 58
Omphalocyclus 10
Oncorhynchus 92
Onoclea sensibilis 235
Onychiopsis 219, 238
elongata 238
mantelli 238, 239
psilotoides 211, 238-239
Onychocella 4, 5, 12, 34
bathonica 3, 4, 5-6, 7, 15, 25, 41
bellona n, 35-36, 41; PI. 6, figs. 5-7
cf. £e£Aa ii, 34-35, 41; PI. 6, figs, i, 2
flabelliformis 6
hagenowi 6
koninckiana 6
lamarcki 35
wys# 34, 41 ; PI. 6, figs. 3, 4
piriformis 5, 6, n, 41; PI. 5, figs, n, 12
Onychocellaria 36
caecilia n, 36-37, 41; PL 5, figs. 1-6
Onychocellidae 13
Orbicyathus 346
Orbitoides 10
Oreosomatidae 96
Orthograptus cf. whitfieldi 189
Orthonotacythere 432
nodosa 432
pulchella 432
sp. 432; PI. 15, fig. i
Osculipora 17
truncata 17, 41
Pachycormidae 98
Pachypeteris 240
lancsolata 240-241 *-242 ; PI. i, fig. 6
Pagruo 87
Palaeocentrotus 81, 97
Palaehystrichophora infusorioides 109
cf. paucisetosa 109
Palaeoperidinium castanea 131
spinosum 142, 143
ventriosum 130, 131
Palaeostomocystis echinulata 157
Paleschara 13, 14
Palescharidae 13
Paracoscinus 363, 364
Paracycas cteis 246
Paracyclotosaurus 292
Paracyprididae 385, 399
Paracypris 399
terraefullonicae 399; PI. 7, figs. 2, 4
Paracytheridea 433
blakei 433; PI. 15, figs. 4, 5, 9, 10
caytonensis 433
Parakannemeyeria 285, 290
Paranacyathus 312, 354
magnipora 311
Parariscus 433
bathonicus 433; PI. 15, figs. 2, 8
Parasudis 81
Paratrachichthys 68
Pareodiniaceae 130
Pareodina sp. 109
Parupeneus 91
Pateroperca 51-52, 53, 54, 55, 83, 84, 86, 87, 93, 98
libanica 51*, 52, 53
Pavobeisselina 40
Pecopteris reticulata 239
Pegasiformes 96
Pelletieria 232
valdensis 232-233*-234*-235; PI. 6, figs, i, 2
Pellia 220
Peltostega 291
Pentaceropsis 91
recurvirostris 91*, 92
Pentacerotidae 87, 91
Percichthyidae 87, 89, 92
Percidae 87
Perciformes 49, 87, 88, 92, 93, 95, 96, 99
Percoidei 86, 89, 92, 96
Permo-Trias continental sediments classification-
Africa 293
Petalopora sp. 18, 41; PI. i, figs. 1-3
Pharmacichthyidae 63-65, 92, 97, 100
Pharmacichthys 49, 63, 65, 66, 84, 87, 88, 93, 94,
95. 97. 98, 99
venenifer 63
Pinacocyathus 342
spicularis 342, 343*-344; PI. 12, figs, i, 2
Placerias 290
P/a/a# 87
Platycopina 380, 395
Platycythere 431
verriculata 431
sp. 431 ; PI. 14, fig. 5
Plecoglossus 86
Pleurograpsus linearis 186
Pleurograptus linearis 186-187*; PI. 2, figs. 1-3
linearis simplex 186, 187
Pleuronectes 89
Pleuronectiformes 93, 95
Pleuronectoidei 89
Pliophloea 37
gluma 37
sp. 7, 14, 15, 37, 41; PI. 7, fig. 3
Pithodella 24
Podocopida 380, 395
Podocopina 383, 397
Poly cope 427
45°
fungosa 427-428; PI. 16, figs. 4, 9
Polycopodae 427
Polycoscinidae 334
Polycoscinus 334, 369
Polydactylus 87
Polymixia 56, 57, 58, 59, 60, 61, 88, 94, 95
japonicus 57
nobilis 56*, 57
Polymixiidae 56-57, 61, 92, 93, 95, 98
Polymixioidei 49, 56, 63, 66, 84, 86, 88, 99
Polynemoidei 87
" Polyphyma bulbosa "12
Polypodiaceae 235
Poly pr ion 87
Polysphaeridium 151
laminaspinosum 151; PI. 4, figs. 10, n
pumilum 151
Polystichum 235, 236
Polystillicidocyathus 318, 327
Pomacanthus 87
Pomadasyidae 87
Pomatomidae 87
Pomatomus 87
Pontocyprella 385
harrisiana 379, 385; PI. 2, fig. 3
Porina 13
cenomana 13
flabellata 40
Porinidae 13
Praechara symmetrica 215
Praeschuleridea 387, 405
schwageriana 387; PI. 2, fig. 6
subtrigona intermedia 387
magna 405
subtrigona 387, 405; PI. 8, figs. 4-6
sp. 388; PI. 2, fig. 8
Pristolepis 89
Procytheridea 430
ininuta 430
parva 430
Progonocythere 389, 423
blakeana 393
juglandica 389
stilla423; PI. 13, fig. 2
Progonocytheridae 388, 409, 428
Progonocytherinae 388, 409
Prelates 88
Prolixosphaeridium 160, 161
conulum 160-161; PI 8, figs. 5, 6
granulosum 161
mixtispinosum 161
Prosoporella 37
cornuta 37
Protocheilostomata 14
Protocytheridae 430
Protopharetra 348, 354, 356, 370
graphica 346, 348
Psettodes 89
Pseudoceratium turneri 171
INDEX
Pseudocycas dunkeriana 212
Pseudotorellia 248, 250
angustifolia 249
crassifolia 249
ensiformis 249, 250
ephela 249, 250, 251
grojecensis 249, 251
heterophylla 248-250*-251 ; PI. 6, figs. 6, 7
longifolia 249
minuta 249, 250, 251
nordenskioldi 248, 249, 250
Pteridophyta 210, 211, 224
Pteridospermae 210, 211, 240
Pterophyllum schaumburgense 242
Puesto Viejo Formation 290
Puncturiella 33
cf. superba 33-34, 41; PI. 4, figs. 8-10
Pustulopora benedeniana 16
Pycnoidocoscinus 363, 367, 368
pycnoideum 367-368; PI. 18, figs. 4-8
Pycnosterinx 59-60, 99
discoides 59
dubius 59, 60, 61, 88
gracilis 59
/afws 59, 60
russeggerii 59*, 60
Pycnosteroides 65, 66, 67, 68, 84, 87, 88, 93, 95, 99
levispinosus 65, 66*
Pycnosteroididae 65-66, 92, 98
Pyripora texana 13
Quadricellaria 29
elegans 29, 41; PI. 6, figs. 8-10
excavata 29
pulchella 29 ,
Radulopora 3, 21-22
minor 3, 21, 22-23, 31, 41; PI. 3, figs. 6-10
radula 21, 22*; PI. 3, figs, n, 12
Rastrites 198
capillaris 198, 199*
carnicus 201
distans 200
gemmatus 198
linnaei 200, 201*
maximus 184, 199, 200-201*-202; PI. 5, fig. 7
cf. peregrinus 200
spina 199
Rectocythere 409
lincolnensis 410
rugosa 409
sugillata 409-410; PI. 10, figs. 3-6
Rectonychocella 27
Red Marl (Beds) 258, 259, 292, 293
Regularia 302, 312, 346, 370
Reptescharinella 29, 30
Retecava abbottii 16
lichenoides 18
ramosa 16
INDEX
Retecoscinus 325, 367
Retepora lichenoides 18
truncata 17
Reteporidae 18
lichenoides 15, 18, 41
Reticrisina 12
" Reticulipora dianthus "12
Retropinna 86
Rhabdocnema 302
Rhabdocyathus 302, 304
tubexternus 304
Rhabdolynthus 302
Rhagasostoma bellona 35
simplex 30
Rhammatopora gaultina 1 3
(?) johnstoniana 13
pembrokiae 13
tn'wei 13
Rhiniopora 8
jurassica 7
scabra 7, 8
Rhytidosteus 291
Robustocyathidae 311
Robustocyathus 311, 312, 369
ar genius 311
annulatus 311
artecaveatus 311
biohermicus 311
dewsws 311
hupei 311
levigatus 311
magnipora 311-312; PI. 3, fig. 5
moori 311
novus 311
polyseptatus 311
proskurjakowi 311
pseudotichus 311, 314
salebrosus 311
spinosus 311
stapipora 309
subacutus 311, 312-313*-314, 369; PI. 3, figs.
6,7
sucharichensis 311
tennis 311
Rosseliana 31
thomasi 3, 7, n, 14, 19, 31, 41; PI. 4, figs. 4-6
Ruffordia 229
goepperti 229-230*-231-232; PI. 4, figs. 1-7;
PL 5, figs. 1-4
var. latifolia 229
Sajanocyathus 369
Salairocyathidae 332, 333, 336
Salairocyathus 336
(Polystillicidocyathus) 336
erbosimilis 336
(Salairocyathus) 336
annulatus 336, 337*-338, 369, 372 ; PI. n, fig. 4
pospelovi 336
zenkovae 336
Salmo 84
Salmoniformes 81
Sangusaurus 285, 286, 289, 290, 292
Sangusaurus edentatus 257, 286-287*-288*-
289*-290
Santa Maria Formation 281, 283, 284, 290
Sardinioides 82, 83, 98
attenuatus 83, 98
Saurida 81, 89
Scalenodontoides 291
Schedophilus 87
Schizaeaceae 229, 231, 232, 235
Schuleridea 385, 402, 403
(Eoschuleridea) 402
bathonica 386, 387, 403
horatiana 402-404. [403*] ; PI. 7, fig. 7
subperforata 386; PI. 2, fig. 4
trigonalis 386-387, 388; PI. 2, fig. 7
(Schuleridea) jonesiana 379, 385 ; PI. 2, fig. 5
Schulerideidae 385, 402
Schulerideinae 385, 402
Sciaena 87
Sciaenidae 87
Scolopsis 87
Scorpaemiformes 95, 96
Scorpididae 92
Selaginella 224, 227
dawsoni 224-225-226 *-227
dichotoma 227
emmeliana 227
Aa//ei 224, 227
sanguinolenta 227
Selaginellaceae 224
Selaginellites 224
dawsoni 224
nosikovii 227
polaris 227
Selene 98
Semieschara complanata 31
Semiescharinella 29, 30
complanata n, 29-30, 41 ; PI. 4, figs. 11-15
Semiescharipora cornuta 37
Semiophorus 97
Serranidae 87, 92
Sertulariadae 192
Siderolites 10
Siniperca 63, 89
chuatsi go*
Sinokannemeyeria 285, 290
Siphonella elegans 20
Smittipora 27
Soleoidei 89
Sparidae 87
Sparus 87
Spermatophyta 240
Sphaerodon 87
Sphenocephalidae 61-62, 92, 98
Sphenocephalus 92, 93, 94, 95, 99
452
fissicaudus 61, 62*
kurriana 212
sternbergiana 212
Sphenopteris fittoni 238
fontainei 238
mantelli 238
ruff or di 238
Sphyraena 87, 88, 92
Sphyraenoidei 87
Spinosocyathus 342
Spirillicyathus 344 ^
Spirocyathella 344
Spirocyathus atlanticus 348
irregularis 344, 348
major 349
speciosus 350
Spirograptus 197
intermedius 197
planus 197
Stahleckeria 257, 282, 283, 284
potens 282*
Stahleckeriidae 281
Stamenocella 25
inferaviculifera 25
marginata 6, n, 25-26, 41; PI. 3, figs. 1-5
mediaviculifera 25
Stephanoberycoidei 68
Stichocentrus 77-78, 94, 95
liratus 77, 78*
Stichopora regularis 16
Stillicidocyathidae 317, 319
Stillicidocyathus 318
Stromateoidei 87
Stomatopora smithi 9
Surculosphaeridium no, 158
longifurcatum 158-159; PI. 8, fig. 9
Synbranchiformes 96
Synodontidae 82
Sy nodus 81
foetens 57
Syringocnema 301, 370, 371
Tabellaecyathidae 370
Tabellaecyathus 348
Taeniopteris beyrichii var. superba 244
Teilhardia valdensis 238
Tannuolacyathidae 325
Tannuolacyathus 325
Tanyosphaeridium 151, 160
variecalamum 151; PI. 6, figs. 2, 5
Tarpon 54, 84, 85
Terquemula 393, 423
blakeana 393, 423; PI. 4, fig. 8; PI. 5, fig. i
Tetraodontiformes 95
Tetrasphaera 132
Thalamocyathidae 317
Thalamocyathus 317, 318, 319
flexuosus 318
trachealis 318, 319
INDEX
Thallites 219, 220-221
catenelloides 222*, 223-224
uralensis 223
valdensis 221-222*-223
willsi 224
yaiei 220
zeilleri 219
Thallophyta 210, 211, 214
Thamnograptus 185
Thecidea 40
papillata n, 40, 41, 42; PI. i, figs. 9, 10
Thrissops 84, 85
Thyracella 26
cf. meudonensis 26, 41; PI. 2, figs. 4-6
Tomocyathus 334, 340, 369, 372
(Erugatocyathus) 299, 334, 372
compositus 334
echinus 334
£*'»* 334
kundatus 334
papillatus 334-335*-336; PI. n, fig. 2
shoriensis 334
(Tomocyathus} michniaki 334
operosus 334
Tortula 218
Trachichthyidae 67, 68, 69, 77, 94, 95, 96
Trachichthyoides ornatus 78
Trachichthys 68
Trachinocephalus 81
Trachycythere 428
verrucosa 428
sp. 428; PI. 1 6, fig. 3
Trachyleberididae 394, 401, 428
Trachyleberidinae 401
Trichodinium 131, 132, 162
castaneum 131-132; PI. u, figs, i, 2, 3
intermedium 132
paucispinum 163
Trichodium castanea 131
Triletes 224
Truncatula tetrasticha 17
Truncatula truncata 17
Tselfatiidae 98
Tubantia 74
cataphractus 74*
Tumuliolynthus 302, 304, 369
irregularis 304-305; PI. i, fig. i
karakolensis 304
musatovi 304
tubexternus 304
vologdini 304
Tumulocoscininae 338
Tumulocoscinus 338
Tunkia 302
Tuvacyathus 325
f/mo karooensis 259
INDEX
453
Velifer Si, 97
Veliferidae 78, 98
Velumella 27
Vincularia 27, 28
canalifera 27-28, 41; PI. 5, figs. 7-10; PI.
figs. 9-12
concinna 28; PI. 8, fig. 13
flexuosa 27, 28; PI. 8, fig. 14
fragilis 27
rugica 32
transversa 32
undata 32
Volvacyathus 354, 356, 370
Vomer 98
Vomeridae 98
Vomeropsis 98
Weichselia 236, 239
mantelli 236, 239
reticulata 236, 239; PI. 6, fig. 5
W etlugosaurus 291
Wilbertopora mutabilis 13
Worthenopora 13, 14
Worthenoporidae 13
Xanthidium ramosum 172
spinosum 166
tubiferum 143
complex 146
Xenichthys 87
Xenistitius 87
Xenocys 87
Xenolepidichthys 96
Zambiasaurus 257, 259, 260-261, 281, 282, 283,
284, 287, 290, 292
submersus 257, 262-263*-264*-265-266*-
267-268*-269*-270*-271*-272*-273*-
274*-275*-276*-277*-278*-279*-280*-
281*-282*-286
Zeidae 96
Zeiformes 49, 93, 96, 97, 100
Zews 96
Zonacyathus 314-315, 369
flexum 369
poletaevae 369
retevallum 314, 315-316, 317, PI. 4, figs. 4, 5
retezona 315, 316*-317; PI. 4, figs. 1-3
vermiculatum 369
Zone of Climacograptus wilsoni 189, 190, 195
Dicranograptus clingani 185, 188, 192, 193, 195
Didymograptus extensus 195
Monograptus convolutus 196
Monograptus cyphus 193
Monograptus gregarius 196
Monograptus riccartonensis 203
Monograptus sedgwickii 196
Orthograptus vesiculosus 193
Pleurograptus linearis 185, 188, 192, 193, 195
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