SS ee 20

~ : :

Bulletin of the foeneives

British Museum (Natural eelies)

Cenomanian brachiopods from the
Lower Chalk of Britain and
northern Europe

E. F. Owen

Geology series Vol44 No2 28 July 1988

The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific
series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an
Historical series.

Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum’s resources. Many of the papers are
works of reference that will remain indispensable for years to come.

Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of the
series on either an Annual or Per Volume basis. Prices vary according to the contents of the
individual parts. Orders and enquiries should be sent to:

Publications Sales,
British Museum (Natural History),
Cromwell Road,

London SW7 5BD,
bias England.
an
SH MUS EAS
Ain ea DN
, i I tas “tig, ¥4 \
ieee ae \ ;
Nene eter dl
SEG NUE Ss serie ten iS Lule ‘
Be : eo

nll f

i

World List abbreviation: Bull. Br. Mus. nat. Hist. (Geol.) i
‘

£

© British Museum (Natural History), 1988 ‘
4

The Geology Series is edited in the Museum’s Department of Palaeontology 4
Keeper of Palaeontology: Dr L.R.M. Cocks hy
Editor of the Bulletin: Dr M. K. Howarth |
Assistant Editor: Mr D. L. F. Sealy 4
if

ISBN 0 56507022 3
ISSN 0007-1471 i
Geology series %
British Museum (Natural History) Vol 44 No 2 pp 65-175
Cromwell Road q
London SW7 SBD Issued 28 July 1988 3

Cenomanian brachiopods from the Lower Chalk of:
Britain and northern Europe

E. F. Owen

Department of Palaeontology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

Contents
SWMODSS coaccovscondmeacadeoou tae Cae te er erect o Acie Oren eae es a aia aa 67
Introduction................ cis pe CCG Ae A OC OAR TERE Laer 67
Distnbution-ofsome Cenomanian rocks’ peete-ceece sees ee seeeeee ee eee ees eee ce cs 69
‘SWISS GIES Sn S010) 1 onpencer Cad HG ae nGBE.cOnSh SAG AABROne PSS AnGuSosane Soe noo Craoben Renee 81
arvihy Ibin ni Woke IM Sl <r aren mene entacntn eenaaee HERE m ant coc re amr ar ara natant: aaa 83
GenuswlingulaBru cule nacre ace ecace cies cacti hee ee ee Oe: 83
ingularsubovalis: Davidsonespeti reece eee ee eee 83
amilvaNhynchonellidaeGrraye.c<.s85 coc ast ska 259515 4 + soles Ho oo ele er pee nalean erie 84
Subfamily, @yclothyrininae Maknidinyecseeceee eee eee eee eee ree ae ee 84
GenusiGyclothirisuMs Coyivaget nec cce soso nesoaense ah acre ein ee eat seas aes 84
Cyclothyris difformis (Valenciennes, in Lamarck) ...................00ecceeeeeeeee 84
GyclothyrisypunfieldensisispanOVaneasssssces tos eee race etna meee er 85
(CIVEOU AS [OARS OSINONS, sepeeecacepybetbbbees sade Gacnded od saan oBpesaanoAnoaaeE 85
GyclothyrisjulgnetispynOVeraecese oe seeee rere eene ce ee eee Ren stgeee ents 86
Cyclothyris compressa (Valenciennes, in Lamarck) ..................0eeccete cece ee 86
Gyclothyrisilamarckianal dOxrbigny) encase eee eee eae eee 89
GenuswBunninnynchiaiOWeMls sos eee ea et as Shot deers 89
BUTninnyNGRiatdevoniana Sp iMOVA= Aerie eee a one eae eee eee 89
Sul familyaW NCEntalMye rani eens eves revereclnanertre ais tro Tae Aree Alen a nate HRN Se NI dn divin 90
GenuskGrasinhynGhiai© wens casa sate ec ss hos cess eck or ene ae ce eee 90
Grasinhynchia\grastana (d:Orbigny)) eas. .2 see ec ae seen ee ene ase eee ene elon 90
Grasirhynchiamartini(Mantell)\ eee a--ceeace cee aeee eo eens eee eee 91
Family Wellerellidae Licharev, in Rzhonsnitskaya ..................ccceee eee eeeeeneeees 92
GenusiO;biriynchiaikettittaeerence scaesene eee cece oceans eer 92
Orbirhynchia mantelliana (J. de C. Sowerby) ..........65..ccc ccc ve cece cece eeeeees 92
Orbiujuynchiawiesti(Quenstedt)ieee-seeeee eee eee eee eer eee cee eee 98
O;binhnynchiamulticostataRettitteecn mace esecens een ee eee eee 99
Orbirnynchiaiwilmingtonensis'spsnOvs ceeereeee kee ee eee ae ee eee eee eee 102
OrbighynchiaiboussensisispynOv> Seecncceweos se soto Seen oe es wo ae 103
pamilweN Onellid aevA Ben, ee trea as savas. aegnniae denise Ghietaea meet schee on caeuuldabile aioe 103
SubfamilysMonticlarellinae)@hildsieeecenens ss cee ee eee era eters 103
GenuseMionticlarellaWASMiewsk ayer ccc rate tpecers «ore rele oe aie eee oie ele eee 103
Monticlarellaibrepinostisi(ROCMEN) | reerreeecre ere ote eee beeee ene ccneereecie: 103
Hanaiilypherebratulid ae Gray) apse cs narroce oct dece ee cine sais ae ae alee cle rele acineRtose/slorsineterneiniss 104
Subfamily Sellithyridinae Muir-Wood ............... 0c see e cece eee e cece eee eeeeeeeeeneeee 104
GenusisellithyrissMiddlemissenc-creccnoececrtcn tices nee: seer aaa 104
Sellithyrisitornacensis|(dvArchiac)i- pees: eee ee ae ee hoe eee eas dee ee ee 104
Sellithyris phaseolina (Valenciennes, in Lamarck) ....................00000000 eee 106
Genus OvatathiyrismOva saree ces acre eee erin ele eee a Laciaides Bia tle vlales ae hetaleteietece hele 106
Ovatathyrisiovata\(Ss Sowerby) foeeeeceee sce ei eisi-t ctesseiscecereaceacd acess ocean oe 107
Ovatathynis potternensiSispemOVe wee ctr ecre aimee coe ieieteceereie cise eee veer ies 109
GenusiBoubeithyrisi\€oxeSaMiddlemiss seer. e ce teie eee etetetereis o> eisai siete wale cle ee »_ Wil
Boubeithyris boubei (d’Archiac) ................0eeece scence teen eee eee ee eee ees 112
Boubeithyris diploplicata sp. NOV. ..........:.0cceeeeee nee n nee ee ene eee ee teeta ees 112

Bull. Br. Mus. nat. Hist. (Geol.) 44 (2): 65-175 Issued 28 July 1988

66 E. F. OWEN

Subfamily Rectithyridinae Muir-Wood ...................0c0eccceecccee eee eeeeeeeeeeees 114
Genus) Rectithyrisisahnierereeeesat eee eee cree eee eee eee eee eee eer 114
IRectithynisswrightonum)spanOve ee cece eEEe eee eeeeeeee rece cce eee eee ee 114
(Genus#inopeothynisi SMilnOVAeeee ee eee Eee eee eee eer eee ener eerie eee ECC eEee 116
liropeothyniswvectisispanoNeucasceet eae ee cee eee eee eee eee eee eee ce 116

Genus) MoutonithyrissMiddlenissieeepeee reece reeeeee ee eeee eee eee eee ee eeree 122
Moutonithyris dutempleana (d’Orbigny) ............-....22-.0e eee cece cere eee eee 122
WMOLEORELASITES CAG|HO 50) WON: cn onanrc000e6005000000060000000cvv0bseesacasceonoCUNba: 126
SubfamilygNenthebrochinael|€oopemrree cere ee eee eee eee eee een eee eee eee eet ree 127
Genustiiarmatosial Coopeteen oe aeeeeee eee eee eee eee eee eee eee sah sansa ena 127
Harmatosiaicrassa dvATrchiac)eaereee cece ree ee eee aero eee ee eee 127
PamilyiGibbithynididae! Muir Wood seerrereree eee eae e terre errr eee rere errr attr 128
Subfamily Gibbithyridinae Muir-Wood ................0.cc cece cece ce ence eee nneeeeees 128
Genus) Concinnithyris!Sahn 2 ace ctercaacen see sone ee sere eeeeie see eee eee 128
Concinnithyrisiobesal(ndel@sSowerby)i-eereeer ere e ee eee eee eee ee eee erEr 128
Concinnithyris subundata (J. Sowerby) .............--+000 eee eee eee cee ee eens 132

Genus OrnatothyrissSaln oo... sisters siccices os «oie cioisses si etepateislas= eevee te etasion eee ae 134
Ornatothyrisisulcifera (Mons) ieepeeee eee eee eee eee eee eee eee eee 138
Subfamulys;€apillithyridinae|€oopemeresreeeee eee a eee eee esate eee ee er eee 142
Genus! Capillithyris\ Katz. <r cates greetcon sc ouisjutasore nes sae os eel pee chee ee CCE EEE EE 142
Gapillithyrisisquamosa| (Mantel) Reeeeseece eee eee eee eee eee eee eee 142
Family ?Boreiothyridae Dagys .................. ccc cece cence eect e een ennneneeneees 144
Genus Arcuatothyris Popiel-Barczyk .............. 00.000 cece eee e eee eee eee 144
Arcuatothyris arcuata (ROeMET) ......... 2.62 eee cece eee eee eee tenn ees 144
Hamily/Cancellothyrididaeihomsontereereeee eee eee eee EEC EEE ee eee eee 147
Subfamily Cancellothyridinae Thomson .................---.0e eee eee eee e eect ee eee sees 147
GenuswierebratulinadOrbignyseesasecee-eee eee eee eee eee eee EEE Eee eee eee 147
Terebratulina etheridgei nom. NOV. .................00ceee cece eect nent eee e eee e ees 147
dierebratulinamodulos@Etherid geass ee eee eee eee eee eee 147
HherebratulinanmbricataispynOVereereeer rece cere en cere eee eee eee Eee eee eee 148
MerebratulinayprovostriatulaispanOVeee Eee eee ree erRe re eee Eee ce ere eerereee 150
Fiamily Zeillerndae: Allany tases. ocjaeetosess sto cde eee io eee eiies tae 154
Genus Modestella Owen, in Casey ..............c0ccccee cece eee tn eee eneceeneeenneenae 154
Modestella geinitzi (Schloenbach) ..................cceee cece eee e cece cte eee e scenes 154
Family Mesathyididae Dall s...3.57 sac eeeeena ser aen carpi aern seine eee ee ier ee eee 156
Genus Argyrotheca Dall. 3. saNeewsdoencetenswoonitaeee eee ae eee nee eRe 156
“Argyrotheca’ megatrema (J. de C. Sowerby) .............--.0e cece cece eee tence eens 156
Family Dallinidae:Beechen «052-2. ..csyeriereserejareniaotere sisters tot eleanor are ePtre earn eee 158
Subfamily (Uncertain eet. ccccacdc wasenctresaseh ae eee oo RCE eee 158
Genus Arenaciarcula Elliott. j.ccc.cccce seed saec ae eee One eee eee eee nee eee ea eee 158
Arenaciarcula beaumonti (d’Archiac) ............0.. cc ee ccc eee c eect ence eee een eeees 158
Hamily Terebratellidae Kane fesse jsateceaee ces cornet ce ena oes besa GG ee een ne eee eee 158
SubfamulysirigonoseminaesElliottite--eeeee cere eeeeer eee rere ee eee eee r ee tener rere 158
Genus eretasBlliott: ccc fa hope Tee ote eres eee OE are lo EE OE 158
Deretaipectita (SU; Sowerby) eas peesen ce eoct ataaee erence aera eer reenee corer eerne 158
Dereta pectita ballardensis subsp. nov. ..............00c.ceecceee cece ee eeneeeeues 162
Deretaiincerta (Davidson)! saccsse anoscote ace COE Eee Ge eRe ce eee 165
Family Thecideidae' Gray. seco cco aa ae ea ee ee er eee Oo NSE 165
Subfamily Mhecideinae! Gray soccer sc ook aCe Oe eee 165
Genus Thecidiopsis|Ochlertq-- are eee eee ence are ee eee 165
dihecidiopsisjessensis| (INOemen) ere eeeeeeeee CEE Eee eee eee eee eee e eee er een enere 165
Flaunallidistribution®. 5st eae roe TIPS fe tote eset tes nc LO ecco Ucar 166
Acknowledgements.) 28. cee soeec ce een ee ECC e ee OEE eee 167
Referencesycicsiccitiacn eeepc nba. 2 Oe eee eRe CET EE CECE cee 168

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 67

Synopsis

A brief history of Cretaceous brachiopod research over the past 120 years is followed by an account of
the distribution of the major brachiopod-bearing outcrops of Lower Chalk (Cenomanian) age in Britain
and northern Europe. The apparent control that some substrates had in the distribution of faunal
assemblages is described, and also their relationship to changes in lithology. The more important Ceno-
manian brachiopod taxa are described and updated, and several new stratigraphically important taxa are
proposed, including the new terebratulacean genus Ovatathyris.

New species of Rhynchonellaceae are: Cyclothyris punfieldensis, C. formosa, C. juigneti, Burrirhynchia
devonia, Orbirhynchia wilmingtonensis and O. boussensis. New species of Terebratulaceae are: Ovatathyris
potternensis, Boubeithyris diploplicata, Rectithyris wrightorum, Tropeothyris vectis, Moutonithyris anglia,
Terebratulina imbricata and T. protostriatula, the latter pro the ‘Terebratula striatula Mantell’ of authors,
non Mantell. Terebratulina etheridgei nom. nov. is proposed to replace T. triangularis Etheridge, non Tate.
The new subspecies Dereta pectita ballardensis is also proposed.

Lectotypes are selected of Lingula subovalis Davidson, Sellithyris phaseolina (Valenciennes, in Lamarck)
and Concinnithyris subundata (J. Sowerby). Neotypes are designated for Sellithyris tornacensis (d’Archiac)
and Terebratulina nodulosa Etheridge.

Introduction

There are few major works on Cretaceous brachiopods beyond those of northern Europe and
the United Kingdom and most of the important works have been confined to comparatively
sparse local faunas. The following account, therefore, deals mainly with British, French and
German species, except for a few forms described from outside this area.

It is important to note that, while the work here is chiefly concerned with species of Cenoma-
nian age, it has been necessary to include a brief mention of some Albian species. These are, in
the main, long-ranging forms, such as those from the Upper Greensand, or facies-controlled
species of Cenomanian aspect such as occur in the Shenley Limestone, Leighton Buzzard,
Bedfordshire, which are of undoubted Lower Albian age.

Great Britain

In Britain many Cretaceous brachiopods were named and described for the first time by James
Sowerby (1815-18), and later his son, James de Carle Sowerby (1826-45), usually under the
single generic name Terebratula. Cenomanian species described by the Sowerbys and con-
sidered to be valid include T. lyra, T. pectita, T. ovata, T. obesa, T. mantelliana, T. obtusa, T.
subundata and T. megatrema. These and several other species have been revised here and
provide most of the systematic part of this paper.

At the time that the Sowerbys were writing, very little information on stratigraphical age was
available, and many specimens described and figured by them were accompanied by only a
brief lithological description of the beds from which they came. Many of their species were
based upon young stages of growth and, moreover, it is in some cases impossible to trace
original material, especially since the Sowerby Collection, now in the British Museum (Natural
History), is far from complete. Subsequent authors have often identified specimens by compari-
son with the Sowerby published illustrations alone and, consequently, many species which they
have described are not always accurately recorded.

The work of the Sowerbys was largely superseded by the publication of Thomas Davidson’s
monographs (1852-54, 1874), which showed that he had a very profound comprehension of the
phylum generally. His contribution to brachiopod taxonomy as a whole is still highly regarded
and most of the Cretaceous species which he described are currently accepted.

Several decades elapsed before further work of significance was produced on British Creta-
ceous brachiopods. Sahni (1929) began a work on British Chalk Terebratulidae with discussion
of previous research; he concluded ‘. . . it is doubtful if any value except historical attaches to
the works cited above’. The works of Davidson and d’Orbigny were among those especially
selected for critical discussion. Sahni ends his comments with the rather startling announce-
ment that ‘the confusion which has obtained in the nomenclature of the Chalk terebratulid
species for over a hundred years has now been cleared.’ It is a pity that Sahni was so critical,

68 E. F. OWEN

because his own work is based almost entirely on poorly documented museum collections. He
created a vast number of new names, largely based upon minor morphological variations. His
attitude to the palaeontological opinions of previous workers contrasts strangely with his
acceptance of museum labels as a key to stratigraphy, and his horizons are, at best, very vague.
Where definite horizons are given, they are often very inaccurate, e.g. Terebratula abrupta Tate
from the Upper Campanian of Northern Ireland is cited as Cenomanian. In fact it is very
difficult to discuss Sahni’s work, as any criticism is bound to be destructive. Nevertheless, his
work does represent the first attempt to separate Cretaceous Terebratulidae into generic
groups.

Pettitt’s (1949, 1954) monograph on Chalk Rhynchonellidae dealt largely with species from
Turonian to Lower Maastrichtian. This was an advance on the work of both Davidson and
d’Orbigny in that it separated the two main generic groups in the Upper Cretaceous, represent-
ed by the new genera Cretirhynchia and Orbirhynchia. Pettitt’s work was not complete and is
not fully representative of the Chalk Rhynchonellidae, especially those of the Lower Chalk.

The most intensive study of the Cretaceous Terebratulidae in recent years has been by F. A.
Middlemiss. His researches have concentrated largely on Lower Cretaceous species, although
he has recently referred the Belgian Tourtia species Terebratula tornacensis d’Archiac to his
genus Sellithyris, originally described from the Aptian (Middlemiss 1959). Middlemess has also
shown that several genera and species thought to have been confined to the Lower Cretaceous
in fact range up into the Upper Cretaceous.

A monographic study of the Terebratulacea from the Trias to Recent by G. A. Cooper (1983)
includes a splendid series of hand-dissected specimens exposing brachidia and cardinalia. This
unique work has greatly advanced our knowledge of the relationship of Cretaceous genera and
species.

During the course of my own research, I have dealt with several genera which have similarly
long stratigraphical ranges and appear well established in both Aptian and Cenomanian beds.
One of these is the rhynchonellid genus Cyclothyris (Owen 1962). The dallinid Kingena David-
son was also reviewed (Owen 1970), and was shown to have a stratigraphical range from the
Lower Albian to Upper Campanian, with two species in the Lower Chalk. Since then, a paper
describing the morphological similarities in the internal structure of the Terebratellaceae was
published (Owen 1977), in which possible evolutionary links between some genera and species
were discussed.

France

The French Cretaceous was intensively studied towards the middle and end of the nineteenth
century and many stratigraphical works appeared, which include original descriptions of
brachiopod species. Brachiopods were described by Valenciennes in Lamarck’s (1819) mono-
graph of the fossils of France, which was partly contemporaneous with the work of the
Sowerbys. Some of the taxa proposed by Valenciennes lacked stratigraphical details and were
poorly illustrated. Among those which can be applied correctly to Cenomanian species are
Terebratula menardi, T. phaseolina, T. difformis and T. compressa. Many points of confusion
were cleared up by Davidson (1850), who revised and figured many of Valenciennes’ species.

D’Archiac’s (1847) description of the Belgian Tourtia species was followed by d’Orbigny’s
monumental work (1848-51). This included chapters on stratigraphical sequences and led to a
scheme for the classification of the Cretaceous strata as a whole, which was published later in
more detail (d’Orbigny 1852). In fact it is to d’Orbigny that we owe the stage name Cenoma-
nian, it having been derived from the Roman name for Le Mans (Sarthe) where the beds are
exposed.

The significance of brachiopod faunas in beds of the Lower Chalk was quickly realized by
stratigraphers and many lists appeared of the species described by d’Orbigny. Barrois (1875,
1876) and Bigot (1893) were among those who acknowledged the importance of brachiopods at
this stratigraphical level, especially where ammonites were scarce or absent. Recent contribu-
tions to the stratigraphy of Normandy and the Sarthe include the works of Kennedy &
Hancock (1970), Juignet, Kennedy & Wright (1973), and Juignet & Kennedy (1976).

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 69

Germany

Early in the nineteenth century there was also considerable interest in the Cretaceous of
Germany. Early German monographs included those of Schlotheim (1813, 1820) in which many
Upper Cretaceous brachiopods were illustrated, including some of Cenomanian age. Later,
Roemer (1840) described the brachiopod faunas of the Hilsconglomerat and the Essen Green-
sand.

Additional German species were erected by Quenstedt (1868-71), Geinitz (1872), Dames
(1874), Diecke (1878) and Tiessen (1895). The one name which stands paramount among
German workers, however, is Schloenbach (1867), whose concise and detailed descriptions were
enhanced by good illustrations.

The early interest in the Cretaceous brachiopods was not maintained and, in keeping with
most countries in western Europe, no notable works devoted to Cenomanian brachiopods have
appeared from Germany since before the first World War.

Denmark

Although the Cenomanian is poorly exposed in Denmark, Ravn (1916, 1925) monographed the
fauna from the Arnager Greensand of Bornholm. Two species of Terebratella, T. kofoedi and T.
bornholmensis, were described as new and a third species was identified as Terebratella beau-
monti (d’Archiac) (Ravn 1916). This species was referred to Arenaciarcula in the revision by
Owen (1977: 236).

Distribution of some Cenomanian rocks

The Weald

Although there are many accounts of the geology of the Weald ranging from the works of
Topley (1875) to the more detailed account of Jukes-Browne & Hill (1896, 1900-04), perhaps
the most detailed descriptions of the Chalk of Sussex are those of Gaster (1929, 1937, 1939,
1944, 1951). He visited numerous localities where the Lower Chalk was well exposed in
working quarries. Few are still available, but Gaster recorded brachiopods from most sections
he visited, and his collection, presented to the British Museum (Natural History) in 1957, has
been used during the course of the present study.

Classic sections, such as those recorded by Mantell (1822) in the Lewes district, have long
been overgrown and lost, but there are still some exposures in large working quarries southeast
of Lewes which present good sections through most of the Cenomanian. Some of these sections
were cited by Kennedy (1969), whose work has been closely followed here, and many of the
brachiopods he collected are used in the present study. Among the many exposures described
by Kennedy are the pits for the cement industry in the Southerham and Glynde areas. East-
wood’s Cement Co. pit, Southerham Grey Pit, Newington’s pit, Rodmill Cement Works pit
near Asham and the Balcombe pit are among those which have yielded many fine brachiopods.
As a result of his work on the coastal sections at Folkestone and Eastbourne, Kennedy was
able to apply ammonite Zones to sections in the pits (Table 1).

The coastal sections, however, are complicated by numerous faults and superficial slumping,
and there is no clear section from Eastbourne towards Beachy Head. South of Cow Gap, the
2-3m thick Glauconitic Marl can be traced in bluffs at the base of the cliffs to Head Ledge,
where it is well exposed and contains brachiopods with well-preserved shells, including the
Grasirhynchia grasiana (d’Orbigny), Monticlarella rectifrons (Pictet) and Kingena concinna
Owen assemblage.

Approximately 12m above the basement bed at Eastbourne, a bed containing Orbirhynchia
mantelliana (J. de C. Sowerby) in abundance forms a recognizable band. It can be better seen
where the beds are at beach level on a platform east of Beachy Head. This is band 6 of
Kennedy (1969), which occurs at the top of the Turrilites costatus Subzone, Acanthoceras
rhotomagense Zone.

Higher up, at the top of band 9 of Kennedy (Acanthoceras jukesbrownei Zone) Concinnithyris
subundata (J. de C. Sowerby) occurs with a marked variant burhamensis Sahni, but specimens of

70

E. F. OWEN

Fig. 1 Sketch-map of the major outcrops of the
Lower Chalk in England. Only the major
towns in each area are marked.

Cambridge

Warminster

Salisbury
Chard

° 2‘
Wilmington rR
andown
CE Compto! ‘ 4
Swanage Bay (0) Miles 50

this variety collected here are proportionately smaller than similarly-named forms from the
type locality at Burham, Kent.

During the course of a more detailed study of coastal and inland areas of Sussex for a Ph.D.
thesis (not completed), Helen Anderson measured sections previously studied by Kennedy. In
some cases she reached significantly different results, and she identified two distinct
Orbirhynchia mantelliana bands: see p. 93. This confirms the unpublished results obtained by
C. J. Wood, British Geological Survey (personal communication), from the Channel Tunnel no.

1 (Aycliffe) and Channel Tunnel no. 2 (Dover Pier) boreholes near Dover, and the Fetcham
Mill borehole, Leatherhead, Surrey.

71

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE

——————————————

1gXDs
Sp4ao1jjaqUuD W

1)]2]UDUL
sp4aoyjaiuv W

4osanzavad
spAad0yIUDIp

asuabpwojoy4
SpAa0YyIUDIp

wunjsngo4

sp4a2094 DD
WNSSDAI

sp4ad094 0D

a4pjnoiapu
spsad0IA]DD

Sajs4adns
pisabv 4

wnup1juvd
Sp4ao]a1UD W

1u0x1p

‘18 $p.4991]]91UD A

asuabowojoy-
SpAaI0YyUDIp

laumoagsaynt
spsa20yluvIp

aupjno1apu
Spsa20IA DD

wnupiuysab

‘13 Sp4220910]aW

asuaopv40jor
SDABIOUIID AA

aSuaUD]LIADI

sp4a20buyisoa
HG PSENS

sp4ao1]a1UuD W

SnjvjSoo

saqy14in
snjnov

saqyidan J,

yjaqubu
Sp4aJaqUuD

1uoxIp
sp4aoyjaiuv W

asuabowuojoya
spsaz0yjuDoy

laumougsaynt
spaaz0yIuDoy

14aBuvsanb
SPABIOIA“IDD

wnuniuysab
$1429091013 JW

ippnt
SDAIIOIPADIOI NN

asuaopv.10jo09
SDABIOUIID AA

snuiqjn
sa11joqiyoa Ny

yjajupu
Sp4aI1]a1UD IW

adUuoZ asuabowojoy

auo0Z snuajd

ajouizia
$p42914N]D]

yjaquput
sp4aI1]a1UD WV

1uojbuiuund
spsaoojpyduony

1yjvds
spsa20oavIn|_

19Z]0]
$a]1aZzJOT

ldajjanu
$D4189091010W
wnuviuysab

$D499091010 W

lB Uh NON =

Sa ld dl ae el GIN
a a NVINVINONAO—

YaMOT
‘NOUNL

(S961 [ewoyL)
aouely “| ‘S

(9L61 ‘1D 12 oxpoury)

steuuojnog

souozqns

SOUOZ

(p861 Apouusy 2 14311)
UIBILIG

(6L61 IPruYyoS 7% Isusq)
AuBulies) WION

(6L6] UueUIpelM)

Auvulies) 410K

a SS eee eee eee

‘odoing WJ9yjJOU pue UreILIg JO URIURUIOUAD ay} J0J saWIaYyos [RUZ

EC LAB

V2 E. F. OWEN

Inland exposures along the North Downs

The general succession remains much the same as that seen in the coastal section at Folkestone,
although exposures of Glauconitic Marl are not so readily available in this area. The Chalk
Marl is well represented with the transition to the more massive Grey Chalk rather better
defined than on the coast. The best localities are in the Medway Valley where a series of
quarries were described by Dibley (1918) and Kennedy (1969). The brachiopod faunas are in a
confused state of nomenclature and are in need of revision. The present work, in addition to
being a re-appraisal of Kennedy’s work, is aimed at updating and redescribing some of the
brachiopod species thought to be of stratigraphical importance. These are, in the main, listed
by Kennedy (1969: 482, 485, 486) in his description of the beds exposed at Bluebell Hill,
Burham, Kent.

Hampshire
Better known for its Upper Chalk and Tertiary deposits, Hampshire also includes geologically
the Isle of Wight, where the Upper Greensand and Lower Chalk are well exposed in the cliff

Compton Bay Gore Cliff & Rocken End Culver Cliff

lens of chert’

P B
Ophosphatized & gsog’ bes

unphosphatized H.
carcitanensis assem-
blage ammonites
(Kennedy, 1969)
®phosphatized
Cenomanian
ammonites
@phosphatized S.

fe=marly chalk
dispar Zone

. |=: Phosphates
ammonites
(Wright & Wright,
1942) ™

ji x abundant sponges
U. Greensand
2cs with stone bands
ig with chert layers
== GAULT CLAY

Fig. 2 Correlation of the Upper Greensand and Cenomanian beds exposed at Compton Bay, Gore
Cliff and Culver Cliff, Isle of Wight.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 13}

sections from Compton Bay in the west to Culver cliff in the east, Fig. 2. The brachiopod fauna
is again represented in bands in two distinct facies, the marl beds of the Middle Cenomanian,
containing the same or a similar assemblage of Grasirhynchia grasiana, G. martini, Kingena
concinna, Terebratulina protostriatula sp. nov. (p. 150), and Orbirhynchia mantelliana, as on the
mainland. The more sandy facies of the Upper Greensand and Glauconitic Marl have a
brachiopod fauna similar to those at Eastbourne and the south-western localities in the
Warminster and south Devon areas. As in the case of the inland exposures of the Weald, some
elements of the fauna appear to be relatively independent of facies and also occur in beds of a
more sandy and glauconitic nature, notably Grasirhynchia grasiana and Terebratulina proto-
striatula sp. nov. The genus Cyclothyris, however, appears to be confined to the coarser sedi-
ments, except for Cyclothyris sp. and C. difformis which have been recorded from the chalky
basement beds of Dorset and Wiltshire (Kennedy 1970).

South-western outcrops

Accounts of the Upper Cretaceous of south-west England can be found in the works of
Jukes-Browne & Hill (1903-4), Wilson et al. (1958), Drummond (1970), Kennedy (1970) and
Jarvis & Woodroof (1984).

Dorset. Except at the coast, the brachiopod faunas in Dorset occur mainly in the Chalk
Basement Bed. Brachiopods are predominantly Terebratulidae and have been collected from
Evershot, Eggardon, Chilfrome, Maiden Newton, Beaminster, Askerswell, Toller Fraterum and
Toller Porcorum and other places listed by Kennedy (1970).

Kennedy’s (1970) detailed account of the Cenomanian Basement Beds in Wiltshire and north
Dorset was accompanied by numerous graphic sections of the sequence indicating the ammon-
ite assemblages and brachiopod faunal bands. This has proved invaluable since, hitherto, only
the broadest outlines of horizons and zones were to be found on labels in museums and private
collections.

At the coast, especially in the Punfield, Swanage, area, Fig. 3, there is lateral variation in beds
of Upper Albian to Cenomanian age. In the cliff section west of Ballard Point, the basal 15cm
of the Chalk Marl contains an Orbirhynchia mantelliana band with associated Grasirhynchia

Chalk basement bed -
basal 10-15¢em, full of dark/

e light brown phosphatic pebbles
in a glauconitic marl.

1

Nodules contain phosphates,
Considerable {|| 4 as does intervening glauc-
lateral vari- | onitic sand - highly complex
ation at this Mile unit - some material piped
level, over 1 down from above.
very short j Large, rather smooth surf-
distance. H aced cobbles of light sand-

mt b stone with variable

development.

Nodular sandstone with
abundant oysters.

--------- Break in nodularity.
Glauconite sand with rather
scattered nodules.

fo)

Fig. 3. Vertical section through part of the Upper Albian and Cenomanian at Ballard Point near
Punfield Cove, Swanage Bay, Dorset. The Dereta pectita ballardensis beds occur at the base of the
nodular sandstone with oysters.

74 E. F. OWEN

grasiana and an undescribed Concinnithyris sp. The Cenomanian Basement Bed (Fig. 3e) is of
Middle Cenomanian age, and rests on a highly complex unit (Fig. 3b—d) containing phosphatic
nodules and glauconitic sands showing evidence of piped material from the beds above. It
contains both Upper Albian and Cenomanian ammonites. A layer of limestone boulders forms
a convenient marker separating the Cenomanian beds from the underlying Albian beds (Fig.
3a) with numerous Amphidonte obliquata (Pulteney). Just beneath these boulders is a bed
containing examples of Dereta pectita that differ from the typical form described by J. Sowerby
(1816) from Horningsham, Wiltshire, in their subquadrate outline, larger foramen and coarser
ornament of radiating costae. This distinctive form is described here as Dereta pectita ballard-
ensis subsp. nov. (p. 162).

Devon. The farthest western outcrop of Cenomanian rocks occurs in the Haldon Hills
(Hamblin & Wood 1976; Wood in Selwood et al. 1984), but the succession can be more readily
studied at the coast between Branscombe and Humble Point where this part of it is well

PINNACLES - DEVON ST. 2205677

nodular chalk

TURONIAN

nodular glauconitic

limestone

glauconitic limestone

BED C

glauconitic limestone with basal
layer of abundant phosphates
BED C

intensely bioturbated
glauconitic limestone

BED B

CENOMANIAN

nodular limestone
BED B
nodular limestone with phosphatized

fossils & cobbles
BED A2

Fig. 4 Vertical section through the Pinnacles, Devon, showing the expanded Beds C and B which
are well exposed.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE WS

exposed, although totally different in development. Here the four lithological units described by
Jukes-Browne (1903) as divisions A, A2, B and C, can be examined. They were revised by Jarvis
& Woodroof (1984), who gave new names to the divisions. The oldest, division A (Pounds Pool
and Hooken of Jarvis & Woodroof), is a coarse shaly limestone with glauconite and fragments
of quartz, and is very fossiliferous. Although some of the ammonites in these beds show signs of
reworking, the molluscan and brachiopod faunas are well preserved, with specimens of Cyclo-
thyris difformis (Valenciennes in Lamarck), Dereta pectita (J. Sowerby) and Kingena arenosa

WILMINGTON (WHITE HART) - DEVON SieeZ08n9 99

calcareous greensand with phosphates overlain by nodular chalk

sandy limestone with

prominent piping

GRIZZLE with abundant Holaster spp.

nodular calcareous

sandstone (fossiliferous)

Zi
ssf
Z
<
=
O
7a
Lu
U

calcareous sands

sandstone cobbles in greensand

first layer of chert

cross: stratified sandstone

UPPER GREENSAND

Fig. 5 Section at White Hart sand pit, Wilmington, Devon, showing the extent of the calcareous
sands and the Grizzle above. See also Smith et al. (1988: 8).

76 E. F. OWEN

(d’Archiac). According to Kennedy (1970) the age is Mantelliceras saxbii Subzone, M. mantelli
Zone.

Bed B (Little Beach of Jarvis & Woodroof) is a finer, highly glauconitic, white sandy lime-
stone containing no brachiopods but having ammonites of the Mantelliceras dixoni Zone.

Bed C (Pinnacles of Jarvis & Woodroof) is developed on the coast as a hard, gritty, highly
glauconitic nodular chalk, streaked with red and green veins and highly burrowed with Tha-
lassinoides paradoxica. There are three ammonite assemblages: phosphatized jukesbrownei and
guerangeri Zone elements, glauconitized geslinianum elements and indigenous ? geslinianum
Zone elements, and numerous bivalves, serpulids and bryozoans. The brachiopod fauna con-
sists of Orbirhynchia wiesti (Quenstedt), O. multicostata Pettitt, Concinnithyris sp. and
Arcuatothyris rugulosa (Morris). This is a similar brachiopod fauna to that of the Plenus Marl
elsewhere.

W. E. Smith (1957a, b, 1961) described the lateral variations in the lithology and thickness of
the coastal outcrops and some of their inland equivalents, particularly those exposed in the two
sandpits in the Wilmington district further north. These pits, Hutchin’s sandpit (ST 216003) and
the more accessible White Hart sand pit (SY 208999), Fig. 5, were reviewed by Kennedy (1970:
66, 665). Carter & Hart (1977: 99, 100) reviewed the work on the basis of planktonic micro-
faunal evidence, but their findings do not alter the information regarding the position of the
brachiopod-bearing beds. Jarvis & Woodroof (1984) have now given what might be a more
acceptable explanation of the age and position of the beds in this section. The lithostratigraphy
of the Wilmington section and its correlation with the coastal section have recently been fully
documented by A. B. Smith et al. (1988).

The Wilmington outlier consists of loose-bedded sands, sandy nodular limestones and glau-
conitic chalk. The nodular limestone or “Grizzle’ contains a rich fauna of molluscs, echinoderms
and brachiopods. Mantelliceras spp., Schloenbachia spp., and Inoceramus virgatus are com-
monly associated with a fauna of predominantly rhynchonellid brachiopods. Cyclothyris dif-
formis is a particularly common species, occurring with Dereta pectita, Terebrirostra lyra,
Arenaciarcula beaumonti, Monticlarella brevirostris, Orbirhynchia wilmingtonensis sp. nov., Ova-
tathyris ovata and Kingena arenosa. An example of the north German species Gemmarcula
canaliculata, which also occurs at Wilmington and Tournai, Belgium, is figured here, Pl. 17, figs
19-21.

Eastern outcrops
The Cambridge Greensand was formerly exposed in phosphate workings south and east of the
town. Nowadays, sections are confined to those incidental to chalk workings, as in the Rugby
Cement Co. quarry at Barrington, where it is possible to see the Cambridge Greensand with
the overlying Chalk Marl and Totternhoe Stone. The age of the Cambridge Greensand has
been much discussed: Carter & Hart (1977), from microfaunal evidence, believe that it belongs
to the Neostlingoceras carcitanense Subzone, at the base of the Lower Cenomanian. Brachio-
pods occur chiefly at the base of the Cambridge Greensand. Many are crushed, decorticated
and phosphatized, and are believed to have been derived from the Upper Albian (Morter &
Wood 1983). In the Fulbourne area, where the Lower Chalk is well exposed, specimens of
Ornatothyris sulcifera (Morris) have been found in the top 2-3m of the section. The full
geological range of this species, however, is not known.

The Plenus Marls are represented in the Cambridge area, but are reduced in thickness. The
characteristic rhynchonellid Orbirhynchia multicostata is present in moderate numbers but the
zone fossil, the belemnite Actinocamax plenus, is rare in this district.

Norfolk

The Cretaceous rocks exposed on the Norfolk coast include the Red Rock of Hunstanton Cliff
which rests on the Carstone and represents the Middle to Upper Albian in East Anglia. It
consists of three distinctly different lithologies which have been distinguished by various
authors as Beds A, B & C or their equivalent beds 3, 2 & 1 (Fig. 6). More recently, Gallois &
Morter (1978) have introduced another numerical scheme for these beds which begins at the

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE Wi

Lower. Chalk

CARSTONE =

Fig. 6 The cliff section at Hunstanton, Norfolk, to show Beds A, B, and C of the Red Rock, the
Inoceramus Bed and the Totternhoe Stone.

dentatus Zone of the Middle Albian and ends at Bed 19 at the top of the dispar Zone of the
Upper Albian. Brachiopods occur in Beds 13 to 16 where the fauna is dominated by Mou-
tonithyris dutempleana, Orbirhynchia parkinsoni and Kingena spinulosa, species which are indi-
genous in the basal beds of the Cambridge Greensand, considered to be of Lower Cenomanian
age. These beds are condensed and only a few metres thick at Hunstanton, and correlation with
the much thicker succession in Lincolnshire and Yorkshire is difficult (Morter & Wood 1983).

The hard bands of consolidated limestone known as the ‘Paradoxica’ Bed, chalkstone with a
hardground, and the Inoceramus Beds, marls rich in bioclastic debris, are clearly visible above
the Red Rock at Hunstanton (Fig. 6). They contain a varied brachiopod fauna of Ornatothyris
spp., O. pentagonalis, Concinnithyris sp. and Kingena concinna.

North-east England
The paucity of ammonites in the Cenomanian of Lincolnshire, Humberside and North York-
shire has made direct correlation with beds of equivalent age in southern England very difficult.
Certain well-marked lithological horizons have been recognized, however, and can be equated
with beds in eastern and southern England. Brachiopod faunas in these lithological bands have
become more important since Bower & Farmery (1910), Jefferies (1963) and Jeans (1973)
indicated brachiopod bands which appear to be common to both northern and southern areas.
Bower & Farmery (1910) established a numerical system for the beds in the Lower Chalk of
Lincolnshire which was broadly applicable to the entire northern province. Their Bed I com-
prised the ‘Paradoxica’ Bed and the two Inoceramus Beds. Between these and the Grey Bed
(Totternhoe Stone), a series of hard bluish-grey chalk beds occur, within which a band of

78 E. F. OWEN

Orbirhynchia sp. is well developed; Jeans (1968) called it the Lower Orbirhynchia Bed. Bower &
Farmery attached a great deal of importance to the existence of two bands in their Beds V-IX,
the Lower and Upper Pink Bands occurring above the Totternhoe Stone. The Lower Pink
Band is the more fossiliferous and contains Grasirhynchia martini, Kingena concinna and Tereb-
ratulina ornata (Roemer) (described here as Terebratulina imbricata sp. nov., p. 148). The
presence of large numbers of T. ornata suggested to these authors that the Lower Pink Band
could be called the T. ornata Subzone. Wright & Wright (1942) recorded many Terebratulina
from the Lower Pink Band in Yorkshire.

A burrowed erosion surface at the top of Bed IX is overlain by Bed X, a conglomerate at the
base followed by the laminated marls of the Black Band, both of Neocardioceras juddi Zone,
Upper Cenomanian, age. The conglomerate is highly fossiliferous and contains Monticlarella
Jefferiesi, Orbirhynchia multicostata and Ornatothyris sp., a brachiopod assemblage referred to
by Jefferies (1963) who concluded that it characterized the upper part of the Plenus Marls.

Northern Ireland

Upper Cretaceous sediments outcrop around the margins of the Antrim basalt plateau, but
Cenomanian deposits are restricted to the south-east in Co. Antrim. Here they consist of a
series of sands forming part of the Hibernian Greensands extending from south of Belfast to
Portmuck, north of Island Magee. The stratigraphy has been revised by McGugan (1957), Reid

Portmuck

Woodburn Glen

Carrickfergus

Cave Hill

WindyaGapnl©
ee eee ICTs

Colin Glen

(0) 5 Miles

Fig. 7 Sketch-map showing the major outcrops of Cenomanian rocks in the Belfast area and Co.
Antrim.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 79

(1958a, b, 1971), Hancock (1961), Fletcher (1977) and Fletcher & Wood (1983). The sediments
consist of two main stratigraphical units, the Hibernian Greensand (Cenomanian to Middle
Santonian) and the Ulster White Limestone (Santonian to Maastrichtian). The Hibernian
Greensand includes the Belfast Marls (a new name by Fletcher & Wood, 1982, for the Glauco-
nite Sands) which are of Mantelliceras mantelli Zone age. This unit yields brachiopods in the
Island Magee area, including Cyclothyris sp., Orbirhynchia sp., Capillithyris squamosa and
indeterminate Terebratulidae. The brachiopod fauna has never been adequately investigated
and is in need of revision.

The succeeding Island Magee Siltstone Member (Fletcher & Wood 1982) consists of grey
argillaceous sandstones with hard calcareous bands, and is of upper Acanthoceras rhotoma-
gense Zone age. It is followed non-sequentially by the Colinwell Sands, in which brachiopods
are fairly common, but are not identifiable, and are again in need of revision.

Northern France

At Cap Blanc Nez in the Boulonnais the succession is much the same as at Dover and
Folkestone, and the Lower Chalk, although somewhat more compact, contains an identical
brachiopod fauna. Amedro et al. (1976, 1978) divided this part of the succession into ten parts,
with a letter and a numeral, C—L and 1-27, to denote its position in the top of the Albian and

za

Boulogne x Tournai

Eastbourne Bins

Etretat

at

Le Havre Rouen xBeauvais
a
Villers
: é 4 x Paris
Vimoutiers "J
rn

Troyes «x yd

St.Calais
J
a
¥ xTours ae

Le Mans

ee eM
0 50 100

‘Fig.8 The distribution of the major outcrops of Lower Chalk in NW France.

80 E. F. OWEN

the Cenomanian. Numerous specimens of Orbirhynchia mantelliana occur in Bed H towards the
top of Band 19. This is near the top of the T. costatus Subzone and corresponds exactly to the
two O. mantelliana Bands of southern England discussed earlier. I have collected specimens of
O. mantelliana from a similar band at Cran d’Escalles and, from a bed just below, examples of
Kingena concinna, Modestella geinitzi, Grasirhynchia grasiana, G. martini and Capillithyris squa-
mosa (E. F. & M. M. Owen collection, British Museum (Natural History)).

The classic coastal section at St Jouin, reviewed by Juignet & Kennedy (1976) is typical of
the coastal exposures in Normandy. Lithological variations include cherty bands with sponge
spicules, hard limestones, and bands of marl and sandy limestones. Kingena arenosa, though
somewhat smaller than the typical form from the Tourtia of Belgium, occurs in the Lower and
Middle Cenomanian limestones associated with Terebrirostra lyra and Dereta pectita which are
also smaller than in England. At Octeville and Cap de la Héve, where the glauconite is more
concentrated, there is a predominance of Rhynchonellidae. Here, as in Dorset and Devon,
Cyclothyris difformis is a comparatively common fossil occurring with other brachiopods in the
coarser sandy facies.

The Cretaceous beds in the Rouen area are affected by the Vernon—Rouen—Pavilly anticlinal
axis, and the basal Cenomanian is seen only at Rouen where it consists of bioturbated glauco-
nitic sands resting on grey clays of the Upper Albian. Above this basement bed are 20m of the
Craie Glauconieuse with bands of black chert and hardgrounds, of Mantelliceras mantelli Zone
age, and containing specimens of Cyclothyris cf. difformis and Grasirhynchia grasiana. Above
the Craie Glauconieuse are 2m of Craie de Rouen, of Acanthoceras rhotomagense Zone age,
and in the chalk above the large terebratulid Concinnithyris obesa (J. de C. Sowerby) occurs in
association with Arcuatothyris rugulosa, Capillithyris squamosa disparilis (d’Orbigny) and
Orbirhynchia cf. paucicostata (Roemer).

The chalky facies of Rouen can be traced eastwards through Orne, where it interdigitates
with the sandy facies at Le Mans (Sarthe). Here the fauna changes once more; Cyclothyris
compressa (Valenciennes in Lamarck) characterizes the Upper Cenomanian Sables du Perche,
where it occurs with Gemmarcula menardi (Lamarck), an example of which is figured here
(Pl. 17, figs 4-6), Sellithyris tornacensis (d’Archiac) and S. phaseolina (Lamarck). Similar
assemblages occur at Briollay, Mayet and Meziéres-sous-Ballon.

At Bousse, the fine-grained chalk of the Sables de Bousse contains Sellithyris phaseolina
(Lamarck) and Orbirhynchia boussensis sp. nov. (p. 103). In the Charente, where the sediments
are fine compact chalk, Sellithyris phaseolina occurs, in association with Gemmarcula caranton-
ensis (Pl. 17, figs 22—24).

East of the Paris Basin, a few exposures of Cenomanian age yield brachiopods. At Vitry
(Marne) Concinnithyris obesa occurs rarely in the chalky facies, and at Brienne-le-Chateau
(Aube) an Orbirhynchia sp. similar to O. mantelliana (J. de C. Sowerby) has been found.

Belgium

The brachiopods in the Cenomanian Tourtia deposits at Tournai and Gussignies are similar in
some respects to the faunas from Wilmington, Devon, the Essen Greensand and the Minster
Basin. However, Rhynchonellidae are predominant at Wilmington, while Terebratulidae and
Terebratellidae are predominent at the Belgian localities and at Essen.

Most of the Cenomanian Tourtia occurs around Tournai and Mons and has a very distinc-
tive and varied brachiopod fauna. The Rhynchonellidae include Cyclothyris scaldisensis
(d’Archiac) and Orbirhynchia cf. paucicostata (Roemer), while the Terebratulidae are dominated
by Rectithyris depressa (Valenciennes in Lamarck), Capillithyris capillata (d’Archiac) and Sel-
lithyris tornacensis (d’Archiac). The Terebratellidae include Arenaciarcula beaumonti (d’Archiac)
and Gemmarcula canaliculata (Roemer) (see Pl. 17, figs 22-24), which also occur at the north
German localities, while A. beaumonti occurs also in Poland and at Wilmington, Devon.

Germany
In the Munster Basin, the Cenomanian occurs as many different facies which include arena-
ceous, calcareous and glauconitic marl developments. The more fossiliferous outcrops are

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 81

Munster Basin

e oo
Munster
Essen a
Bochim A

bees
bepes Magdeburg
Saltzgitter? +
ess

cme SCck eae
Dortmund ntils
Dresden
B ~a @
ag,
R b

Paris Boege oo
Basin P ad

A

TOKE Czechoslovakia

Fig. 9 The major Cenomanian outcrops of north Germany, and the Dresden and Regensburg
districts.

mainly in the southern part of the basin in the Essen Greensand where the fauna is similar to
that in the Belgian Tourtia. Additional species not known from the Belgian Tourtia are Bur-
rirhynchia sigma (Schloenbach), Arcuatothyris arcuata (Roemer) and a large undescribed tereb-
ratulid, similar to Concinnithyris davidsoni from the Upper Greensand of Warminster.

Further east at Floteberg (Tecklenberg), Langelsheim and Wunstorf in the Middle Cenoma-
nian marl facies, the familiar Grasirhynchia grasiana, Concinnithyris cf. subundata, Kingena
concinna assemblage occurs, although the lithology is a harder, more compact marly limestone
than occurs in France or southern England.

A fauna similar to that of the Grés du Mans in the Sarthe occurs in the Bryozoensandstein at
Morasheimer. Lehner (1937) listed Gemmarcula menardi, Sellithyris phaseolina and Cyclothyris
compressa. The Upper Cenomanian is also represented at the Kassenberg, Miilheim, where a
species of Orbirhynchia occurs that is similar to O. boussensis sp. nov. of the Sables de Bousse,
Sarthe.

In the basal conglomerate of the Actinocamax plenus Zone at Ratsteinbruch, Dresden, large
Cyclothyris sudetica (Geinitz) occur in pockets of softer rock. The same species also occurs in
the Lower Sandstone at Regensberg, Bavaria. Associated faunas include undescribed Terebra-
tulidae, probably of the genus Sellithyris.

Systematic descriptions

The present research was never intended to be taken as a comprehensive monograph of all
known species of brachiopods in the Lower Chalk, since many of these have been adequately

82

E. F. OWEN

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 83

described or revised in recent years and are in little need of additional remarks. Problematic
subjects have been investigated and the results and their interpretations fully explained. The
extensive use of transverse serial sections, revealing the internal structures of Chalk Terebratu-
lidae, has somewhat altered the established or more conventional interpretation, based largely
on dissected specimens, of M. R. Sahni (1929).

Full synonymies are of limited value where in the cited works there is no reliable horizon
correlation and no convincing illustration accompanying a description of a species and, there-
fore, such items are excluded from the synonymies in the present work.

This section deals with existing genera and species (some with new descriptions) and
describes new taxa where these are considered to be stratigraphically important. Unless other-
wise stated, the material described or referred to is in the collections of the Department of
Palaeontology, British Museum (Natural History), where specimen numbers are normally pre-
fixed B or BB, but reference to material from other collections is denoted by the following
prefixes:

SM = Sedgwick Museum, Cambridge

BGS = British Geological Survey (formerly Institute of Geological Sciences)

GM = Museum d’Histoire Naturelle, Geneva.

In the figures of serial sections the numbers shown by each section give the distance in mm
from the umbo to the first section, thereafter the distance between sections. In the plate
descriptions, SEM = scanning electron micrograph.

The following terms are used: Outline—the complete shell outline viewed dorsally or ven-
trally. Profile—the complete shell outline viewed from either side, or from the front. View(ed)—
the surfaces seen in the described aspect, e.g. ‘anterior view’ includes details of the commissure.
In describing ribbing the terms ‘costa’, ‘costella’ etc. are used in the sense of prominence or
frequency of ribs, not in relation to their origins.

Class INARTICULATA Huxley, 1869
Superfamily LINGULACEA Menke, 1828
Family LINGULIDAE Menke, 1828
Genus LINGULA Bruguicre, 1797

Lingula subovalis Davidson
Pl. 5, fig. 32

1852 Lingula subovalis Davidson: 7; pl. 1, figs 29, 30.

LecToTyYPe. B8291, here selected from three syntypes in the Davidson Collection. Dimensions:
length 12-6mm, width 7-5 mm.

PLATE 1

Figs 1-9 Cyclothyris punfieldensis sp. nov. Figs 1-3, Glauconitic Marl, Undercliff, Isle of Wight.
Paratype BB82244, x 14. Figs. 4-6, Upper Albian, Dispar Zone, Punfield Cove, Dorset. Holotype
BB82245, x 13. Figs 7-9, Basal Cenomanian, Cap de la Héve, Normandy. BB82183, x 15.

Figs 10-12 Cyclothyris cf. punfieldensis sp. nov. Bottom Bed of Upper Albian, Cap de la Heve,
Normandy. BB82182, x 13.

Figs 13-15 Cyclothyris difformis (Valenciennes, in Lamarck). Upper Greensand, Compton Bay, Isle
of Wight. BB82339, x 14.

Figs 16-18 Cyclothyris sp. Albian, Castellas, Narbonne, SW France. BB82367, x 1.

Figs 19-21? Cyclothyris sp. Lower Cenomanian, Chardstock, Devon. B8218, x 2.

Figs 22-24 Burrirhynchia devoniana sp. nov. Lower Cenomanian, White Hart sand pit, Wilmington,
Devon. Holotype BB82252, x 2.

Figs 25-27 Burrirhynchia cf. sigma (Schloenbach). Lower Cenomanian, Annopol, Poland. BB82340,
>< D,

Figs 28-30 Burrirhynchia leightonensis (Walker). Lower Albian, Shenley Hill, Leighton Buzzard,
Bedfordshire. B26575, x 1.

84 E. F. OWEN

EMENDED DESCRIPTION. The outline is regularly elliptical, oval, with subparallel lateral margins;
the valve being slightly convex with an umbonal carina not extending more than one third of
its length. The surface is smooth, marked only by numerous concentric growth-lines.

REMARKS. Living species of Lingula have a habitat restricted to burrows, usually on sandy or
silty shorelines, and sometimes near river estuaries. Their tolerance to depth and temperature is
limited and, if it can be assumed that their habitats to-day are comparable to those of fossils,
lingulas can provide useful evidence of climate and depth.

Apart from the Craniidae, families of inarticulate brachiopods are rare throughout the Meso-
zoic. Species of Lingula have been recorded from the Lower Cretaceous Speeton Clay from time
to time and also occur, though somewhat sporadically, within the Lower Greensand in
southern England and the Isle of Wight. Lingula truncata (J. de C. Sowerby) has been collected
from the Upper Aptian of Sandown and specimens from the Hythe Beds of Kent are in the
Sedgwick Museum, Cambridge. In France a species described by d’Orbigny (1847) as L. raulin-
iana, from probable Upper Aptian clays of Grandpré (Meuse), is possibly a synonym of
Sowerby’s L. truncata.

Records from the Albian are sparse but include a specimen identified by Lang (1904) from
the ‘Gault’ of Black Ven, near Lyme Regis, Dorset. Donovan (1949) recorded Lingula subovalis
from the Albian of Geographical Society Island, Greenland, but this is a misidentification.

There are more records from the Cenomanian, most of which have been identified as Lingula
subovalis or Lingula sp. and include specimens from Butser (Hampshire), Halton, and the
Burwell Rock of Burwell (Cambridgeshire). The youngest specimen recorded from the Cenoma-
nian was identified and figured as Lingula sp. from the Plenus Subzone, Bed 1, at Merstham,
Surrey, by Jefferies (1962).

Class ARTICULATA Huxley, 1869
Superfamily RHYNCHONELLACEA Schuchert, 1900
Family RHYNCHONELLIDAE Gray, 1848
Subfamily CYCLOTHYRININAE Makridin, 1955
Genus CYCLOTHYRIS M‘Coy, 1844

Cyclothyris difformis (Valenciennes, in Lamarck)
Pl. 1, figs 13-15; Pl. 2, figs 4-6, 10-15

1819 Terebratula difformis Valenciennes, in Lamarck: 255; indic. Encyclopedie Methodique (Tabl. Vers,
Coquilles, Mollusques et Polypiers 2): pl. 242, figs 5 (1789). Paris.

1821 Terebratula dimidiata J. Sowerby: 138; pl. 277, fig. 5.

1822 Terebratula gallina Brongniart: 8; pl. 9, fig. 2.

1828 Terebratula deformis Lamarck (sic); Defrance: 160; pl. 5, fig. 3.

1849 Rhynchonella compressa (Lamarck); d’Orbigny: 35; pl. 497, figs 1-6.

1850 Terebratula difformis Lamarck; Davidson: 433; pl. 15, fig. 48.

1852 Terebratula compressa Lamarck; Davidson: 80; pl. 11, figs 4 & 5.

1882 Terebratula difformis Lamarck; Quenstedt: 696; pl. 54.

1900 Rhynchonella difformis (Lamarck); Jukes-Browne: 65, figs 41, 42.

1918 Terebratula difformis Valenciennes, in Lamarck; Clerc & Favre: pl. 15, fig. 84.

1962 Cyclothyris difformis (Valenciennes, in Lamarck) Owen: 51; pl. 5, figs 1-7.

LectotyPr. Lamarck Collection, Museum d’Histoire Naturelle, Geneva, No. 48; from the
Lower Cenomanian, Normandy coast; selected by Owen (1962).

REMARKS. This species was redescribed in a general revision of Cyclothyris M‘Coy by Owen
(1962: 51), where a pictograph illustrated thirteen morphological variants of C. difformis
(Valenciennes, in Lamarck). The present work confirms this variation and adds examples from
several different localities. Pl. 2, figs 1-3 represents the acutely triangular form from the Glau-
conitic Marl of Compton Bay, Isle of Wight and figs 4-6 show an example of a rare form from
the Lower Chalk Basement Beds of Snowdon Hill, Chard, Somerset, in which the anterior

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 85

commissure is asymmetrical. In some assemblages specimens with either right or left asymmetry
are found; for example that figured as PI. 2, figs 10-12, from the Lower Cenomanian of High
Melcombe, Dorset, shows an asymmetrical specimen with left side prominent, and with a
greater number of costae and more numerous concentric growth-lines than on the Chard
specimen.

Cyclothyris punfieldensis sp. nov.
Pl. 1, figs 1-12; Pl. 2, figs 16-18

D1aGNnosis. Transversely oval to broadly triangular in outline; adults measure approximately
18-2mm in length, 14-6mm wide and 14mm thick. Umbo slightly produced, beak suberect.
Large circular foramen surrounded by extended deltidial plates. Anterior commissure low,
arcuate. Dorsal fold poorly developed. Ornament of 30-34 strong, angular, deeply incised
radiating costae.

Name. From the type locality of Punfield Cove.

DESCRIPTION. The broadly oval outline, poorly developed concentric growth lines and faintly
sulcate median dorsal fold are characteristic of this species. In lateral profile the shell is strongly
dorsi-bioconvex, with more inflated dorsal umbonal regions. The consistently arcuate commis-
sure shows no sign of the asymmetry so typical of Cyclothyris difformis (Valenciennes, in
Lamarck) and it can be distinguished from this and other species of Cyclothyris in having a
faint anterior sulcation in the dorsal fold of adult specimens.

HotoryPe. C. W. & E. V. Wright Collection, BB82245; from the Upper Albian, Dispar Zone;
Punfield Cove, Dorset. Length 18-5 mm; width 14-9 mm; thickness 14-0 mm. PI. 1, figs 4-6.

OTHER MATERIAL. In addition to the holotype the following is a paratype: BB82244; Glauco-
nitic Marl; Undercliff, Isle of Wight. Length 18-2 mm; width 13-3mm; thickness 12-7mm. PI. 1,
figs 1-3. Two other specimens are in the BM(NH), BB82183 (PI. 1, figs 7-9) and BB82184.
Other examples are in the British Geological Survey, BGS 110218—110225, Cunnington Collec-
tion from the Upper Greensand of Warminster, Wiltshire. A gerontic example BB82341 is
shown in PI. 2, figs 16-18.

REMARKS. It is probable that in Britain this species is confined to the topmost Upper Albian,
although very similar, conspecific examples occur in the topmost Upper Albian and basal beds
of the Cenomanian at Cap de la Héve, Normandy (PI. 1, figs 7-12). The Normandy examples
resemble the British specimens other than in lacking the characteristic slightly produced umbo.

C. punfieldensis resembles Cyclothyris shenleyensis (Walker) from the Lower Albian of Leigh-
ton Buzzard, Bedfordshire but can be distinguished from that species by its proportionately
wider dimensions, deeper brachial sulcation and higher linguiform extension.

Cyclothyris formosa sp. nov.
Pl. 3, figs 1-6

DiAGnosis. Triangular in outline, trilobate. Umbo short, massive; beak slightly incurved. Beak-
ridges sharp, deltidial plates exposed; interarea fairly extensive. Median dorsal fold well devel-
oped. Ventral valve flat to slightly convex, anteriorly sulcate. Anterior commissure with high
uniplication.

Name. ‘Shapely’.

DESCRIPTION. Convexity of the dorsal valve is regular and its fold, which originated posteriorly
at less than one-third of the valve length, becomes increasingly prominent anteriorly to produce
a strong anterior trilobation in the shell. The surface ornament, of 34-36 rounded costae, is not
interrupted by growth-lines. An average of six to eight costae adorn the median fold with a
corresponding number within the pedicle sulcus. Variation within the species is slight, affecting
only the degree of uniplication.

86 E. F. OWEN

Ho.ortyPe. BB82206; from the Middle Cenomanian (Lennier’s Bed 10), St Juin, Normandy.
Length 22:2 mm; width 24-5 mm; thickness 16-7 mm. PI. 3, figs 1-3.

PARATYPES. BB82207 (PI. 3, figs 4-6), BB82300-31.

REMARKS. Cyclothyris formosa can be distinguished from C. difformis (Valenciennes, in
Lamarck) by its trilobate form, short massive umbo, smaller foramen, lack of asymmetrical
development of the anterior commissure and regular biconvexity. It can be distinguished from
C. juigneti sp. nov. (below), C. compressa (Valenciennes, in Lamarck) and Cyclothyris lamarck-
iana (d’Orbigny) by general outline, more strongly developed uniplication, more rounded and
less deeply incised costae and more regular biconvexity.

Cyclothyris juigneti sp. nov.
Pl. 3, figs 16-21

DiaGnosis. Relatively small, triangular in outline. Umbo short, beak suberect, pointed, with
large auriculate foramen. Brachial valve convex, median fold poorly developed. Pedicle valve
weakly convex. Anterior commissure gently uniplicate with low linguiform extension.

Name. In honour of Dr P. Juignet, University of Caen, Normandy.

DESCRIPTION. The umbo, though short, has a moderately extensive interarea bounded by sharp
beak-ridges. Both valves have approximately 23-25 sharply differentiated costae, with four on
the median fold and three in the sulcus. There is no marginal thickening of the valves and
growth-lines are not visible.

Ho.Lortype. BB82208; from the Middle Cenomanian, Turrilites costatus Zone; La Perriere
Orne, France. Length 17-0mm; width 22:2 mm; thickness 9-8 mm. PI. 3, figs 16-18.

PARATYPES. BB82209 (PI. 3, figs 19-21), BB82210-1.

REMARKS. The species can be distinguished from C. compressa, C. difformis and C. lamarckiana
by its consistently smaller dimensions, more deeply incised and fewer costae, flatter valves and
strongly triangular outline. The dorsal median fold shows about the same degree of develop-
ment as in C. compressa but always remained narrower.

Cyclothyris compressa (Valenciennes, in Lamarck)
Pl. 3, figs 22-27

1819 Terebratula compressa Valenciennes, in Lamarck: 256, no. 54.

1850 Terebratula compressa Lamarck; Davidson: 455; pl. 15, fig. 54.

1918 Terebratula compressa Valenciennes in Lamarck; Clerc & Favre: pl. 19, figs 117, 119.
1962 Cyclothyris compressa (Valenciennes in Lamarck) Owen: 54; pl. 5, fig. 8.

REMARKS. This species was redescribed by Owen (1962), at which time it was thought to

PLATE 2

Figs 1-3 Cyclothyris cf. difformis (Valenciennes, in Lamarck). Upper Greensand, Compton Bay, Isle
of Wight. BB82248, x 15.

Figs 4-6, 10-15 Cyclothyris difformis (Valenciennes, in Lamarck). Figs 4-6, Chalk Basement Bed,
Snowdon Hill, Chard, Somerset. BB82242, x 15. Figs 10-12, Lower Cenomanian, High Mel-
combe, Dorset. BB82243, x 14. Figs 13-15, Middle Cenomanian, nodular limestone, Bovey Lane
Pit, Devon. BB61120, x 15.

Figs 7-9 Cyclothyris scaldisensis (d’Archiac). Bed B equivalent, Cenomanian, St Jouin, Normandy
BB82184, x 14.

Figs 16-18 Cyclothyris punfieldensis sp. nov. (gerontic specimen). Upper Albian, Punfield Cove,
Dorset. BB82341, x 1.

Figs 19-21 Cyclothyris sp. Belfast district, Northern Ireland. B8229, x 14.

87

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE

88 E. F. OWEN

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 89

characterize the Sables du Perche of the Upper Cenomanian in the Le Mans district (Sarthe),
France, being distinct from related forms in both the Marnes a Ostrea biauriculata above and
the Sables et Grés du Mans below. I now believe it to range through all three zones and to
display variation in its valve thickening. The marginal thickening seen on valves of specimens
collected from the Sables du Perche had been interpreted as a more generalized thickening of
the valves, but is now thought to have resulted from local preservation, leaving the rest of the
valve very much thinner. Comparison with material collected from other localities within the
three Upper Cenomanian zones confirms the presence of the species in all three.

Specimens from Mayet (Sarthe) (PI. 3, figs 22-27) have slightly more rounded costae, a less
well defined triangular outline and a wider median fold and sulcus than those of C. difformis or
C. juigneti.

Cyclothyris lamarckiana (d’Orbigny)
Pl. 3, figs 7-15

1849 Rhynchonella lamarckiana d@Orbigny: 32; pl. 496, figs 5—13.

REMARKS. Originally described from the Cenomanian of the Le Mans district, the species
characterizes the Marnes a Ostracées of the Briollay area (Sarthe). It is a comparatively large
species with almost equal biconvexity of valves. The costae are strong and angular and average
30 on each valve, with about six anteriorly on the weak fold and about five in the low ventral
sulcus and linguiform extension. There is a tendency to asymmetry of the anterior commissure,
although this is not shown on the specimens figured here.

DIMENSIONS of figured specimens. BB82203 (PI. 3, figs 7-9): length 21-9mm; width 24-9 mm;
thickness 13:-7mm. BB82204 (PI. 3, figs 10-12): length 21-3mm; width 22:5mm; thickness
14-0 mm. BB82205 (PI. 3, figs 13-15): length 22-6 mm; width 25-7 mm; thickness 16-0 mm.

Genus BURRIRHYNCHIA Owen, 1962

Burrirhynchia devoniana sp. nov.
Pl. 1, figs 22-24

DiAGnNosis. Subtriangular to ovate Burrirhynchia, adults approximately 7-5mm long, 6mm
wide and 10mm thick. Strongly biconvex, uniplicate, dorsal fold low; linguiform extension
trapezoidal, high. Shell surface multicostate. Hinge-teeth large: median septum low, no
septalium.

NAME. ‘From Devon’.

Ho.otyre. BB82252; from the Lower Cenomanian; White Hart sand pit, Wilmington, south
Devon (PI. 1, figs 22—24).

OTHER MATERIAL. In addition to the holotype about fifty specimens were studied. Paratypes
are BB82253-9, BB76321, and SM B10289-93 from the type locality.

DESCRIPTION. Rarely specimens of B. devoniana from the White Hart sand pit show a tendency

PLATE 3

Figs 1-6 Cyclothyris formosa sp. nov. Cenomanian, Lennier’s Bed 10, St Jouin, Normandy. Figs
1-3, Holotype BB82206, x 14. Figs 4-6, Paratype BB82207, x 15.
Figs 7-15 Cyclothyris lamarckiana (d’Orbigny). Marnes a Ostrea biauriculata, Briollay, Sarthe. Figs
7-9, BB82203, x 14. Figs 10-12, BB82204, x 13. Figs 13-15, BB82205, x 15. ;
Figs 16-21 Cyclothyris juigneti sp. nov. Cenomanian, T. costatus Zone, La Perriere (Orne). Figs

16-18, Holotype BB82208, x 14. Figs 19-21, Paratype BB82209, x 13.
Figs 22-27 Cyclothyris compressa (Valenciennes, in Lamarck). Upper Cenomanian, naviculare Zone,
Marnes a Ostrea biauriculata, Mayet (Sarthe). Figs 22-24, BB82211, x 15. Figs 25-27, BB82210,

x 15.

90 E. F. OWEN

towards the typical sulciplicate sigmoid commissure of B. sigma (Schloenbach) from north
Germany (see Schloenbach’s drawing, 1867: pl. 23, figs, 5, 6). A broad sulciplication is also a
feature of the type-species, B. leightonensis (Walker), from the Lower Albian of Leighton
Buzzard, Bedfordshire (PI. 1, figs 28-30).

REMARKS. The chief differences between the Cenomanian B. sigma (Schloenbach) (1867: 500; pl.
23, figs 5-7) and B. devoniana is the costation, which in the former is fine and rounded in
contrast to the fewer (20-25 on each valve with 6 in the sulcus) and more deeply incised ribs on
B. devoniana. Variation is similar to that of the German species, but the outline is more
elongate than in the Lower Albian B. leightonensis. It seems probable that the three species, B.
sigma, B. leightonensis and B. devoniana are related and exhibit adaptations to different
environments, manifested chiefly in the coarseness of their costation.

Subfamily Uncertain
Genus GRASIRHYNCHIA Owen, 1968

Grasirhynchia grasiana (d’Orbigny)
Pl. 5, figs 20-23

1849 Rhynchonella grasiana d’Orbigny: 38; pl. 497, figs 7-10.

1852 Rhynchonella grasiana d’Orbigny; Davidson: 96; pl. 12, figs 17—19.

1867 Rhynchonella grasana d’Orbigny (sic); Schloenbach: 496; pl. 2, figs 8, 9.

1878 Rhynchonella grasana d’Orbigny (sic); Deicke: 26, figs 20, 21.

1895 Rhynchonella grasiana d’Orbigny; Tiessen: 459.

1968 Grasirhynchia grasiana (d’Orbigny) Owen: 20; pl. 2, figs 7a, b, c.

1969 Rhynchonella grasiana d’Orbigny; Panow: 578; pl. 110, fig. 1.

1977 Grasirhynchia grasiana (d’Orbigny); Popiel-Barczyk: 36; pl. 3, figs 3-7; text-figs 10, 11.

LEcTOTYPE. Selected (Owen 1968) from 22 specimens in the d’Orbigny Collection, Museum
d’Histoire Naturelle, Paris, No. 6497 in the d’Orbigny catalogue; from the Cenomanian of Le
Havre, Normandy. Length 13-4mm; width 15mm; thickness 10mm.

REMARKS. The species, the type species of Grasirhynchia, was studied and redescribed (Owen
1968) using transverse serial sections of a British specimen from Sussex which had been care-
fully matched with the lectotype. Since then Popiel-Barczyk (1977) has published transverse
sections through two specimens from the Lower Cenomanian of Annopol, Poland, comparable
to those published by me in 1968. Of her figured sections those of 1977: fig. 10 most closely
resemble my concept of the species.

The stratigraphical range of the species appears to be Lower to Middle Cenomanian. It
exhibits a remarkable ecological tolerance and is found in various lithologies from highly
glauconitic sandy limestones, such as the Upper Greensand of Warminster, Wiltshire and the
Lower Cenomanian Limestone of Wilmington, Devon, to the more marly and chalky facies of
the Middle Cenomanian in parts of Dorset, Sussex and Kent.

The species displays two size variants; the smaller and more finely costate specimens appear-
ing to prefer the more highly calcareous and chalky facies of the western localities. Both forms,
however, occur within the same beds. Further eastwards, in Sussex and Kent, larger, more
coarsely costate specimens become more common, particularly within the marly chalk of the
rhotomagense Zone in the coastal localities. Generally, the smaller form with finer costation is
more strongly convex, having a domed dorsal valve and a moderately highly arched uniplicate
fold. A broad shallow ventral sulcus is present anteriorly in all varieties.

The foramen is characteristically large for a small rhynchonellid, and is partially surrounded
by auriculate extensions of the deltidium. This structure is not confined to G. grasiana, being
present on well-preserved specimens of G. martini (PI. 5, figs 24-26) and on some Cyclothyris
specimens.

Geographically the species has a wide distribution, numerous examples having been collected
from Wilmington, Whitecliff, Haldon Hills and Chardstock, Devon and from the Popple Bed at
Mere, Wiltshire, to Melcombe Bingham, Dorset. A large, coarsely costate form has been

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 91

collected from Heytesbury and Chute Farm, Wiltshire. The species is not common on the Isle
of Wight but examples have been obtained from the Glauconitic Marl of Compton Bay, Gore
Cliff and Rocken End, Undercliff and Culver. A fine specimen was collected by Dr W. J.
Kennedy from the Lower Cenomanian, mantelli Zone, carcitanense Subzone, west side of
Woody Bay.

Continental collections have been made from the type locality and St Jouin in Normandy,
from the more marly beds at Cran d’Escalles near Cap Blanc Nez as well as from Rougefort,
Pas de Calais, Octerville and Vimoutier (Orne). One large specimen is said to come from the
Basses Alpes in southern France; some nearly perfect examples were collected by Dr W. J.
Kennedy from the Middle Cenomanian Sables de Lamnay, at a disused roadside pit near
Courgenard, Sarthe.

Good examples resembling the lectotype were collected by Dr R. Marcinowski from the
Lower Cenomanian of Czestochowa and Julianka in Poland. Other material from the Lower
Cenomanian of Annopol is in the Museum Ziemi, Warsaw, Poland. The species occurs very
rarely in beds considered to be of Middle Cenomanian age in Brochterbeck and Wunstorf,
north Germany.

MATERIAL. The following list of British and some French examples in the BM(NH) is given
with dimensions in mm.

Locality length width thickness BM(NH) No.
Warminster 11-7 13-3 10-4 2326
Chute Farm 9-4 10-7 6:8 98057
10-8 11-4 7:3 ss
10:4 11-4 6:6 A
9-2 9-8 6:5 Fs
9:9 10-0 5:8 x
9-1 9-5 79 5
8:7 8-9 77 Fe
Chardstock 9-0 10-8 7:2 B8145
9-6 8-9 77 &
8 4 8:3 6 4 ”
Undercliff, Isle of Wight 11:5 11:9 77. BB82371
9-1 9-7 6°5 BB82372
Rougefort, Pas de Calais 9-0 8:5 6:8 =
13:4 8-9 5:8 =
121 79 6:2 -
75 8-1 49 -

Grasirhynchia martini (Mantell)
Pl. 5, figs 24-26

1822 Terebratula martini Mantell: 31.

1826 Terebratula pisum J. de C. Sowerby: 70; pl. 536, figs 6, 7.

1838 Terebratula pisum Sowerby; von Buch: 148; pl. 15, fig. 18.

1852 Rhynchonella martini (Mantell) Davidson: 94; pl. 12, fig. 15 only.
1867 Rhynchonella martini (Mantell); Schloenbach: 499; pl. 23, fig. 10.
1895 Rhynchonella martini (Mantell); Tiessen: 458.

1968 Grasirhynchia grasiana (d’Orbigny) Owen: 20; pl. 2, figs 7a, b, c.

Lectotype. B61480 in the Sowerby Collection; selected Owen (1968). Chalk Marl; Hamsey,
Sussex (PI. 5, figs 24-26).

REMARKS. This species is included here because confusion has arisen between examples identi-
fied as G. martini and specimens here assigned to Monticlarella brevirostris (Roemer), p. 103,
originally described from the Planer near Sarstedt, north Germany (Roemer 1840). Davidson

92 E. F. OWEN

(1852) figured three specimens (his pl. 12, figs 15, 16a—d) of which one (fig. 15) is a true G.
martini, while another (fig. 16a—c) is an example of Monticlarella brevirostris from the Chalk
detritus of Charing, Kent. The third specimen (fig. 16d) is unrecognized but came from the
Upper Greensand of Chute Farm, Warminster.

Monticlarella brevirostris (Roemer) is thought to be a senior synonym of Rhynchonella
rectifrons Pictet from the Cretaceous (uncertain horizon) of Ste Croix, Vaud, Switzerland and is
to be the subject of a future paper shared with C. J. Wood. In Britain it appears to be of
stratigraphical significance, occurring at a restricted horizon within the Lower Chalk, and has
been found with G. martini at Folkestone, Eastbourne and localities on the Isle of Wight.

Grasirhynchia martini is more ecologically limited than G. grasiana and appears to be
restricted to the marly beds within the upper part of the Lower and the Middle Cenomanian.
Records of its occurrence in the Lower Cenomanian sandy deposits in the Annopol district of
Poland (Popiel-Barczyk 1977) are doubtful since her figured specimens (pl. 4, figs 1-3) may not
be conspecific, although the serially sectioned specimen from Iwanowice (1977: fig. 12) belongs
to Grasirhynchia and the sections are very like the series published earlier (Owen 1968: text-fig.
2) through a specimen from the Chalk Marl of Hamsey, Sussex.

MATERIAL. B25088, Lower Chalk, Heytesbury; B8139, Lower Chalk, Sussex (Davidson
Collection); B8175, Pit 3, Hallington near Louth, Lincolnshire (J. F. Farmery Collection); one
specimen from the Totternhoe Stone equivalent, Melton, Yorkshire (12/29, F. Whitham Collec-
tion, Hull).

Family WELLERELLIDAE Licharevy, in Rzhonsnitskaya, 1959
Genus ORBIRHYNCHIA Pettitt, 1954
TYPE SPECIES. Orbirhynchia orbignyi Pettitt, 1954.

DISCUSSION. Pettitt (1954: 29) erected Orbirhynchia for his species Orbirhynchia orbignyi, from
the Middle Chalk, Labiatus Zone, of Reigate, Surrey. He differentiated several species in the
genus ranging from the Lower to Upper Chalk and attempted to distinguish his type species, O.
orbignyi, from O. cuvieri by comparison of external morphology and type of costation. His
distinctions are somewhat arbitrary, based on minor morphological differences which could be
attributed to variation within the species, and this casts doubt upon the stratigraphical value of
some of his taxa.

In my opinion, O. cuvieri from the Middle Chalk, and O. wiesti (Quenstedt) and O. multi-
costata Pettitt from the Lower Chalk, should be regarded as conspecific, while O. compta Pettitt
and O. orbignyi are also possible synonyms of O. cuvieri.

Another group of Orbirhynchia species which appears to have many morphological charac-
ters in common is represented by O. mantelliana (J. de C. Sowerby) from the Lower Chalk,
Chalk Marl of Hamsey, Sussex. This group can be distinguished from the O. wiesti group by its
more robust appearance, flatter valves, coarser (fewer) costae and trapezoidal to asymmetrical
anterior commissure. It also differs internally from the type species in having a thicker test and
less well developed dental lamellae. I include the species O. obscura Pettitt, from the Lower
Chalk of Folkestone, in synonymy with O. mantelliana as it appears to be nothing more than
an intraspecific variant. O. parva Pettitt, from the Lower Chalk of Cambridge, is probably
another synonym, but is consistently smaller and broader in general outline than the type
specimen of O. mantelliana and may deserve subspecific status. O. dispansa Pettitt, from the
Upper Chalk, appears to belong in the mantelliana group.

Orbirhynchia mantelliana (J. de C. Sowerby)
Figs 10-13; Pl. 4, figs 1-6, 10-12

1826 Terebratula mantelliana J. de C. Sowerby: pl. 537, fig. 5.
1838 Terebratula mantelliana Sow.; von Buch: 154; pl. 15, fig. 26.
1855 Rhynchonella mantelliana J. de C. Sowerby; Davidson: 87; pl. 12, figs 20, 21.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 93

1954 Orbirhynchia mantelliana (J. de C. Sowerby) Pettitt: 31; pl. 3, figs 10a-c.
1954 Orbirhynchia obscura Pettitt: 35; pl. 3, figs la—c, 2a—c.
1969 Orbirhynchia mantelliana (J. de C. Sowerby); Kennedy: pl. 22, figs 5—7.

LECTOTYPE. Sowerby Collection, B61490; from the Lower Chalk of Hamsey, Sussex; selected
Pettitt (1954). Pl. 4, figs 1-3.

DESCRIPTION. The species was adequately described by Pettitt (1954: 31), who gave the average
dimensions as 12 mm long, 13 mm wide and 9 mm thick. The species is subpentagonal in outline
and oval in anterior profile. The brachial valve is moderately convex with a slight but fairly
broad fold anteriorly. The linguiform extension was described as broad with the anterior
margin straight, but variants within the species tend to have an asymmetrical anterior commis-
sure, a feature rarely found in other Orbirhynchia species. There are 12-18 comparatively deeply
incised costae on each valve, with 4 on the fold and 3 in the sulcus.

INTERNAL CHARACTERS. The thickened shell, strong, partially fused dental lamellae and falcifer
crura are shown in serial sections of a topotype by Pettitt (1954: fig. 10).

REMARKS. In spite of his recognition of material from Bedfordshire, Cambridgeshire, Kent,
Surrey and Sussex, Pettitt’s concept of the species appears to have been based largely upon the
type specimen and comparatively few additional examples. The present study takes a somewhat
broader view of the species, including variants from localities other than those already men-
tioned. A simple statistical analysis has been made from random samples obtained from these
localities and a linear regression is presented here, Fig. 11. In addition, a pictograph illustrating
the range of morphological variation is given, Fig. 12.

The species is regarded as an important stratigraphical marker and has been the subject of
discussions by Jeans (1968) and Kennedy (1969). Kennedy emphasized the importance of an
Orbirhynchia mantelliana Band within the rhotomagense Zone of the Middle Cenomanian in the
southeast of England. He fixed the upper limit of his band as corresponding to the boundary
between the Turrilites costatus and T. acutus assemblages, and in his correlation diagram
(1969: fig. 16) he traced it across the Weald from Folkestone in Kent to Asham, near Bed-
dingham, Sussex.

Recently, in an unpublished summary of research on Lower Chalk stratigraphy of the Sussex
coast and inland areas, Helen Anderson (Brighton Polytechnic) has recognized another
Orbirhynchia mantelliana band at Beachy Head, Eastbourne and Southerham Grey Pit, near
Lewes, approximately 4 metres below that of Kennedy. It is not certain whether either of these
bands can be directly equated with the Orbirhynchia Beds of Jeans (1968), but it seems likely

5mm

Fig. 10 Orbirhynchia mantelliana (J. de C. Sowerby). Transverse serial sections through a specimen
(BB82355, Walker colln) from the Lower Chalk, Chalk Marl, of Hamsey, Sussex.

94 E. F. OWEN

WIDTH

Jk=Lectotype

: 5 10 15

LENGTH

Fig. 11 Length x width graph and regression line for 159 specimens of Orbirhynchia mantelliana
(J. de C. Sowerby) from England.

THICKNESS

5 10 15
LENGTH

Fig. 12 Length x thickness graph and regression line for 159 specimens of Orbirhynchia mantelliana
(J. de C. Sowerby) from England.

that the O. mantelliana specimens occurring in a band just above the Grey Bed, and at a level
equivalent to Bed IV of Bower & Farmery (1910) within the H. subglobosus Zone in Lincoln-
shire and Yorkshire, might be the equivalent of the Upper Orbirhynchia mantelliana Band
(Band 6 of Kennedy, 1969) which occurs in Sussex and Kent.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 95

Schloenbach (1867: 494) described Rhynchonella mantellana [sic] from the mantelli and rhoto-
magense Zones of northwestern Germany, placing in synonymy Sowerby’s original Terebratula
mantelliana and Roemer’s (1841) T. paucicosta. Schloenbach’s figured specimen (pl. 23, fig. 11)
shows a flatter form with a more circular outline and only slightly produced beak; his concept
of O. mantelliana resembles only slightly the typical form which was, admittedly, rather poorly
illustrated by Sowerby (1826). His specimen seems closer to Roemer’s T. paucicosta, from the
Essen Greensand, and could even have been a young form of that species. It also resembles a
specimen figured by Panow (1969: 570; pl. 109, fig. 4) and another by Popiel-Barczyck (1977:
30; pl. 1, figs 7-9) called Orbirhynchia mantelliana from the Lower Cenomanian of Poland
which, in addition, have a more broadly arcuate anterior commissure than true O. mantelliana.

Many geographical and ecological variants have now been included within this species,
largely because their separation can be seen to offer no stratigraphical advantage. The species is
listed prominently from the Totternhoe Stone, or just above it, from the type area in Berkshire,
southwards to Wiltshire and Dorset and as far north as Cambridge, Lincolnshire and York-
shire. There are also examples which fall well within the present range of variation from Cran
dEscalles near Cap Blanc Nez, northern France and from Brochterbeck, north Germany.
Specimens which can neither be included here, nor, so far as I can see, within O. paucicosta,
occur in the Tourtia of Tournai, Belgium. This variant has also been collected from beds of a
similar age at Montigny sur Roc, southern France. Specimens figured as Rhynchonella mantel-
liana by Deicke (1878) and Wanderer (1909) are also probably referable to O. paucicosta
(Roemer).

Examples from the Totternhoe Stone at Burwell, Cambridgeshire, while clearly exhibiting all
the typical morphological characters, almost always maintain a robust outline with acutely

>

Fig. 13 Eight specimens showing the range of variation in Orbirhynchia mantelliana, from the Lower
Chalk, rhotomagense Zone, Folkestone, Kent.

96 E. F. OWEN

inflated valves and coarse, deeply incised costae; they are also larger (PI. 4, figs 10-12). It is
probable that this is the variant recorded by Bosworth (1906) in a band just above the Tot-
ternhoe Stone, which he called the Rhynchonella mantelliana Band, because of the abundance of
large specimens of this species associated with ‘Cidaris’ dissimilis and Terebratulina nodulosa.

MATERIAL AND DIMENSIONS. Because of the confusion surrounding this variable species I have
presented both pictographic and statistical data. The following measurements are taken from a
selection of specimens in the British Museum (Natural History) (prefix B or BB) and the British
Geological Survey (others). Specimens in older collections may only have group registration
numbers. Locality and stratigraphical details are quoted from the museum labels, but where
possible checks have been made with colleagues from the B.G.S. Measurements in mm.

Number length width thickness Number length width thickness
Dover, Kent; rhotomagense Zone: Isleham, Cambridge:
B93747 15-9 15-0 10-3 BGS 73064 13-1 12:9 9-9
B93750 13-4 13-6 8-0 73066 9:9 10-0 7-0
BB37477 12:0 11-6 7-1 73067 9-9 10-0 6:5
BB34478 11-0 11-1 8:8 73068 9:8 9-9 69
BB34479 11-1 11-6 8-1 Norton Ferris, Wiltshire:
B22892 10:9 10-8 6:0 : :

Zb 968 15-2 17-0 11-0
B93749 13-1 13-8 8-9 970 12-1 13-1 741
Margett’s Pit, Burham, Kent: 972 13-0 13-9 9:2
B44670 10:9 10:8 5-9 973 13-0 13-9 9-9
B44671 10:8 10-9 Tl 974 12:5 14-3 8-0
Folkestone, Kent; Grey Chalk ae es ne ie

, Kent; Gre alk: Sree
= O5 pi ta 7 6-0 Hitchin, Herts; Totternhoe Stone:
B25102 11-0 12-0 7-5 37017 11-0 11-2 78
B25102 11-8 12:5 6:5 37019 12-8 12-9 8-3
B25102 11-1 10:8 8-1 Swaffam Prior, Cambridge:
B25102 11-5 11-1 8-2 109617 10-0 9-9 7-5
B25102 11-1 10-8 79 109618 13-9 13-5 9-2
B25102 iNile7/ 12-1 8-1
B25102 11:0 11:6 TPA
B25102 11-2 11-3 7-1
B25102 11-0 10-4 7:8
PLATE 4

Figs 1-6, 10-12 Orbirhynchia mantelliana (J. de C. Sowerby). Figs 1-3, Chalk Marl, Hamsey, E.
Sussex. Lectotype B61490, x 14. Figs 4-6, variant from Lower Chalk, rhotomagense Zone, Folk-
estone, Kent. B25105, x 14. Figs 10-12, Totternhoe Stone, Barrington, Cambridge. BB81020,
x 15.

Figs 7-9 Orbirhynchia cf. mantelliana (J. de C. Sowerby) (?paucicosta Roemer). Lower Cenomanian,
Le Mans district, Sarthe. BB82337, x 14.

Figs 13-18 Orbirhynchia wilmingtonensis sp. nov. Figs 13-15, Craie de Rouen, Cote de Ste Cath-
arine, Rouen. BB82338, x 14. Figs 16-18, Lower Cenomanian, mantelli Zone, Wilmington, Devon.
Holotype BB82132, x 14.

Figs 19-21, 31-33 Orbirhynchia boussensis sp. nov. Upper Cenomanian, Sable de Bousse, Bousse
(Sarthe), France. Figs 19-21, Holotype BB82131, x 14. Figs 31-33, Paratype BB61140, x 15.

Figs 22-24 Orbirhynchia sp. Red Rock, Hunstanton, Norfolk. B19904, x 15.

Figs 25-30 Orbirhynchia wiesti (Quenstedt). Figs 25-27, Cenomanian, Bed B, Little Beach, Beer,
Devon. BB7126, x 15. Figs 28-30, Cenomanian Basement Bed, naviculare Zone, Askerswell,
Dorset. BB82097, x 14.

Figs 34-42 Orbirhynchia multicostata Pettitt. Upper Cenomanian. Figs 34-36, Plenus Subzone,
South Ferriby. BB82342, x 15. Figs 37-39, Plenus Marl, Marham, Norfolk. BB82343, x 14. Figs
40-42, Plenus Marl, Burwell, near Louth, Lincolnshire. BB82344, x 15.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 97

98 E. F. OWEN

These and measurements of other specimens from the Sedgwick Museum are included in the
statistical analysis, the plots of which are given in Figs 1 1—12.

Orbirhynchia wiesti (Quenstedt)
Pl. 4, figs 25-30

1871 Terebratula wiesti Quenstedt: 166; pl. 41, fig. 54.

1874 Rhynchonella wiesti (Quenstedt) Davidson: 66; pl. 8, figs 31, a.
1896 Rhynchonella wiesti (Quenstedt); Jukes-Browne & Hill: 147.

1954 Orbirhynchia wiesti (Quenstedt) Pettitt: 37; pl. 3, figs 14a—-c, 18a—c.
1962 Orbirhynchia wiesti (Quenstedt); Jefferies: pl. 77, fig. 32.

1970 Orbirhynchia compta Pettitt; Kennedy: 645.

LECTOTYPE. Selected by Pettitt (1954) from three specimens figured by Quenstedt (1871: pl. 41,
fig. 54); from the Lower Chalk of Chardstock, Devon, and now in the Geologisches Paldont-
ologisches Institut, University of Tiibigen, Germany. Length 14-3mm; width 12:3mm; thick-
ness 10-6 mm.

REMARKS. Pettitt recorded this species from Beer Head, Hooken Cliff and Chardstock in
Devon, a distribution based on twenty specimens from museum collections. He restricted the
species to those with only 28 costae and rejected Davidson’s (1874) broader definition in which
were included specimens with a total of 30 to 32 ribs. Jukes-Browne & Hill (1896) considered
that Davidson’s definition was too narrow: ‘It is a much more variable species than Davidson
seems to have supposed—the average number of ribs seems to be between 24—26, but there are
many forms which have as few as 18, and others which have as many as 30; the former
resemble Rhynchonella mantelliana except that the ribs are not so angular, and the latter come
so near to the broader varieties of R. cuvieri that when placed beside them, they are indistin-
guishable.’

Pettitt (1954: 31) appears to have accepted these remarks in a general sense, but I think that
Jukes-Browne & Hill were actually referring specifically to specimens from Bed 13 of Meyer
(1874) which, they appear to be claiming, were all of one species.

The situation is somewhat analogous to the position of O. mantelliana where the typical
Sowerby definition has been obscured by the numerous varieties added to the species. Before
1954, when Pettitt figured his lectotype of R. wiesti, the species could only have been inter-
preted from the illustrations of Quenstedt (1871: pl. 41, figs 52-54) and the specimen figured by
Davidson (1874: pl. 8, fig. 31). It is not surprising, therefore, that the interpretation of Jukes-
Browne & Hill (1896) should agree with forms from Bed C of the Cenomanian which are now
thought to include variants of both O. wiesti and O. multicostata.

Jefferies (1962), in dividing certain faunal groups into cold water and warm water inhabit-
ants, placed Orbirhynchia multicostata Pettitt into the cold water group and a specimen he
identified as O. wiesti (pl. 77, figs 2a, 2b), belonging to neither the cold water nor the warm
water group, was placed in a miscellaneous group. In his diagram of the standard succession at
Merstham, Surrey, Jefferies (1962: 611) marked the point, Beds 7-8 within what was at that
time the Metoicoceras gourdoni Zone, where O. wiesti commonly occurs. He also indicated a
position, Bed 1, the Metoicoceras geslinianum Zone, where O. multicostata (p. 99) was said to
be common. The relative position of these species in the Actinocamax plenus Subzone was said
by Jefferies to be maintained throughout the entire Anglo-Paris Basin. However, the justifica-
tion for Jefferies’ two zones has been questioned by Kennedy & Hancock (1978, in Rawson et
al.), and Wright & Kennedy (1981) have since suggested that the top part of the Upper
Cenomanian be known as the Neocardioceras juddi Zone so both brachiopod species occur in
this new zone.

If, as Jefferies suggested, O. wiesti lived in a completely different environment and tem-
perature from that of O. multicostata, it is surprising to find Smith (1957a, b, 1961) recording
both species within a comparatively narrow horizon, Bed C of Jukes-Browne & Hill (1896,
1903) on the south Devon coast. This might be explained by the condensed nature of the bed,

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 99

since Wright & Kennedy (1981: 8) regard it as the equivalent of the Plenus Marls and the top
of the Cenomanian.

The species is recorded from the base of Bed 6 at Bovey Lane sand pit by Smith (1961: 117).
The correlation of Bed 6 with Bed C of the coastal section was confirmed by Kennedy (1970:
660), who cited O. multicostata with O. wiesti and Arcuatothyris rugulosa. Specimens collected
from this pit, identified as O. wiesti, vary in their broader and flatter valves, and slightly more
produced umbo. O. wiesti has also been recorded from Bed 13 (Meyer 1874; Jukes-Browne &
Hill 1896), where it is exceedingly rare.

Specimens have been studied in the British Museum (Natural History), including two from
Bed B, Little Beach, Beer, Devon, and six from Bed C, Hooken Cliff (included in series
BB76345—S4). A typical form is in the Davidson Collection (BB12396), and is said to have come
from the Glauconitic Marl of Chardstock. A range of variants came from the Chalk Basement
Bed of Askerswell (BB7127—30).

Orbirhynchia multicostata Pettitt
Pl. 4, figs 34-42

1874 Rhynchonella wiesti Quenstedt; Davidson: 66; pl. 8, figs 31, a, b.
1903 Rhynchonella wiesti Quenst.; Jukes-Browne & Hill: 136, in part.
1954 Orbirhynchia multicostata Pettitt: 38; pl. 3, figs 20a, b.

1962 Orbirhynchia multicostata Pettitt; Jefferies: pl. 77, figs 2a, b.
1963 Orbirhynchia multicostata Pettitt; Jefferies: 7-18.

LectToryPe. Selected by Pettitt (1954) and one of two specimens figured by him (pl. 3, figs
20a, b) from the Upper Cenomanian, Plenus Subzone; Betchworth, Surrey. British Geological
Survey, GSM 92642.

REMARKS. Pettitt (1954: 38) based the description of this species on two rather poorly preserved
syntypes and mentioned a further ten specimens from the Subglobosus and Labiatus Zones and
the Plenus Subzone of the Lower Chalk in Bedfordshire, Buckinghamshire, Cambridgeshire,
Surrey and Sussex.

Since its original description, the species has been widely quoted from the Plenus Subzone in
southeastern and eastern England and from beds considered to be of equivalent age in south
Devon and Dorset. As a result, the interpretation of the species now includes many variants,
which Pettitt might well have regarded as being outside his original concept. These forms range
morphologically from subcircular to broadly oval in outline, with a short suberect beak and
wide arcuate anterior commissure. A prominent concentric growth-line is commonly present at
about half, or just anterior of half, the length of the shell.

In his original description of the Plenus Subzone of the Anglo-Paris Basin, Jefferies (1962)
wrote that O. multicostata was a common fossil in Beds 1a and 1b of the standard succession of
Merstham, Surrey, and also pointed out the apparently wide geographical distribution of this
brachiopod in Britain and France at approximately the same horizon. He interpreted the
spread of O. multicostata, which he thought originated from an early ancestor at Cap d’Antifer,
as influenced by a shallowing of the outer reaches of the basin; this would probably also
account for the rare occurrence of Monticlarella jefferiesi (formerly quoted as “Rhynchonella
lineolata carteri). The rise in temperature during the succeeding deposition in Bed 2, however,
probably resulted in a reversal of the situation, when M. jefferiesi became comparatively
common and O. multicostata exceedingly rare. The association of O. multicostata with the
obviously shallow water brachiopod Lingula sp. appears to confirm the presence of shallow
water conditions at Bed 1 level, as suggested by Jefferies (1962).

This species exhibits a greater tolerance to ecological variation than any Orbirhynchia species
so far investigated. It appears in marls, limestones and chalky facies without significant varia-
tion. Throughout its extraordinary environmental range it maintains the same robust morpho-
logical features which must have inspired its separation from other species of the same genus at

100 E. F. OWEN

ve
ONG

>

Nae

iy

Hi

PLATE 5

Figs 1-6, 13-15 Terebratulina protostriatula sp. nov. Figs 1-3, Chalk Marl, Hamsey, E. Sussex.
Holotype BB6038, x 14. Figs 4-6, Lower Cenomanian, White Hart sand pit, Wilmington, Devon.
Paratype BB82261, x 14. Figs 13-15, Essen Greensand, Essen, north Germany. B35690, x 13.

Figs 7-9 Terebratulina striatula (Mantell). Upper Chalk, Hamsey, E. Sussex. Holotype B457, x 15.
See also Pl. 20, fig. 3.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 101

Fig. 14 Eight specimens showing the range of variation in Orbirhynchia wilmingtonensis sp. nov.,
from the Lower Cenomanian, White Hart sand pit, Wilmington, Devon.

the time of its description, although the specimens chosen for figuring by Pettitt do little to
distinguish it clearly from O. cuvieri or O. wiesti.

Although from the literature this species does not appear to have been as widely distributed
on the Continent as in the United Kingdom, Jefferies (1963) pointed out its value as a marker
fossil in Bed 1 from Cap Blanc Nez and other northern French localities to Cap d’Antifer on

Figs 10-12 Terebratulina ? auriculata (Roemer). Tourtia, Tournai, Belgium. B35491, x 13.

Fig. 16 Terebratulina nodulosa Etheridge. Red Rock, Bed A, Hunstanton, Norfolk. BB82368, x 4.

Fig.17 Terebratulina etheridgei nom. nov. Cambridge Greensand, Cambridge. BB82369, x 4.

Figs 18-19 Thecidiopsis essensis Backhaus. Essen Greensand, Essen, north Germany. B5190, x 10.

Figs 20-23. Grasirhynchia grasiana (d’Orbigny). Cenomanian, Le Havre. Figs 20-22, Lectotype, Coll.
Inst. de Pal., Paris No. 6497, x 2. Fig. 23, specimen showing characteristic auriculation of the
foramen by the extension of the deltidial plates. BB84908, x 6.

Figs 24-26 Grasirhynchia martini (Mantell). Cenomanian, Chalk Marl, Hamsey, E. Sussex. Lecto-
type B61480, x 3.

Fig. 27 ‘Argyrotheca’ megatrema (J. de C. Sowerby). Cambridge Greensand, Cambridge. BB6097,
x 4

Fig. 28 ‘Argyotheca’ sp. Cenomanian, Essen Greensand, Essen, north Germany. B7218, x 4.
Figs 29-31 Monticlarella carteri (Davidson). Cambridge Greensand, Cambridge. BB44427, x 23.
Fig. 32 Lingula subovalis Davidson. Upper Greensand, Potterne, Wiltshire. B8291, x 13.

102 E. F. OWEN

the coast, and it has been collected in beds of the same age at Cernay-en-Dormois east of the
Paris Basin.

It is possible that the Orbirhynchia commonly found in the Sables de Bousse, Sarthe, may be
a geographical variant of this species, but it maintains a uniformity of shape and size, and its
umbonal characters distinguish it from the typical O. multicostata, so I describe it as a new
separate species, O. boussensis (p. 103). Specimens from Bed C at Bovey Lane, Devon closely
resemble O. multicostata and are probably a local variant of that species.

Orbirhynchia wilmingtonensis sp. nov.
Fig. 14; Pl. 4, figs 13-18
1852 Rhynchonella mantelliana Sow.; Davidson: pl. 12, fig. 22.

1903 Rhynchonella mantelliana Sow.; Jukes-Browne & Hill: 129.
Orbirhynchia mantelliana auctt.

DIAGNOsIS. Distinctly subcircular to triangular in outline. Strongly biconvex, coarsely costate.
Umbo slightly produced, deltidium exposed. Anterior profile almost circular, anterior commis-
sure uniplicate with high arch.

NAME. ‘From Wilmington’.

DESCRIPTION. The costae on both valves are triangular in section and deeply incised. They vary
from 10 to 12 in all, with two or three on the almost imperceptible dorsal median fold and with
a corresponding number in the comparatively deep, anteriorly developed broad sulcus.

Variants have a more circular outline, slightly flatter valves and more prominent umbo. The
species is quite distinct from Orbirhynchia mantelliana (J. de C. Sowerby) with which it has been
confused in the past. The new species differs in having fewer and more deeply incised costae,
more triangular outline and stronger biconvexity. It also has a produced umbo exposing the
deltidium, a feature not often seen in other species of the genus. These features also distinguish
it from O. wiesti and O. multicostata, both of which have more numerous and less deeply incised
costae and a less protruding umbo.

HOLotTyPe. BB82132, from the Lower Cenomanian Mantelliceras mantelli Zone; White Hart
sand pit, Wilmington, Devon. Length 11-6mm; width 11-4mm; thickness 9-4mm. PI. 4, figs
16-18.

OTHER MATERIAL. Numerous specimens in the British Museum (Natural History) from the type
locality only. Paratypes are BB82133—43: for dimensions see below. The species is at present
known only from south Devon.

REMARKS. Although the species has been referred to O. mantelliana by various authors, it
compares more readily with a similarly robust form occurring in the Craie de Rouen and
possibly also from the Lower Cenomanian of the Sarthe. The French specimens, however, are
consistently more circular in outline and have a less marked anterior sulcus than the Devon
species.

In its small number of costae O. wilmingtonensis resembles T. paucicostata Roemer from the
Essen Greensand. The new species compares with Roemer’s illustrations (1841: pl. 7, fig. 6), but
his type material is missing, probably destroyed during World War II, and I have never seen a
similar specimen from any locality in the German Cenomanian. There is, therefore, no evidence
for Roemer’s original concept of O. paucicostata. But a specimen figured by Quenstedt (1871:
pl. 41, figs 24-26), from the Planer, Essen, may have been compared with Roemer’s type
specimen and is probably typical of the species. It represents an uncommon species from the
Greensand at Kassenberg and other localities in the north German Cenomanian. It differs from
O. wilmingtonensis in its considerably smaller size, more circular outline and less produced
umbo. Three further specimens from the Essen Greensand, which also might represent
Roemer’s species, are in the Bruckmann Collection, B15468. They are flatter than the specimen
figured by Roemer (1841) and have a closer resemblance to O. mantelliana (J. de C. Sow.).

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 103

DIMENSIONS of paratypes in the BM(NH). Measurements in mm.

Number length width thickness Number length width thickness
BB82133 12-6 12-0 8-5 BB82139 10-5 10-7 7:2
BB82134 14-1 3-2 10-5 BB82140 11-4 10-3 74
BB82135 11-6 11-4 il BB82141 11-6 10-7 9:4
BB82136 12:3 12-0 93 BB82142 11-4 9-8 8:5
BB82137 12:9 11-8 8-9 BB82143 10-6 9-7 6:6
BB82138 11-2 11-4 8-9

Orbirhynchia boussensis sp. nov.
Pl. 4, figs 19-21, 31-33

DIAGNosIs. Comparatively large Orbirhynchia, subcircular to broadly triangular in outline.
Umbo slightly produced, beak suberect, pointed. Foramen smail surrounded by lateral exten-
sions of the deltidium.

NAME. ‘From Bousse’.

DESCRIPTION. The anterior profile is characteristically oval, with an arcuate uniplicate commis-
sure. There is no dorsal fold and the sulcus is confined anteriorly. Adult variation extends only
to minor differences in the width of shells. About 26-28 rounded or subangular costae orna-
ment each valve, commonly with 9 in the sulcus, although up to 12 have been recorded. Some
specimens have a strong growth-line at about half to two-thirds of their valve length.

Ho.otyre. W. J. Kennedy Collection, BB82131; from the Upper Cenomanian, Sables de
Bousse, Sables a Catopygus obtusus; in a disused pit 180 metres west of Cemetery, SW of
Bousse, Sarthe, France (loc. 12 of Jefferies, 1963). Length 15-O0mm; width 15-9mm; thickness
10-2 mm. PI. 4, figs 19-21.

PARATYPES. BB61121—5S9, K. Evans Collection. Some dimensions are given below.

REMARKS. The species is unknown in Britain, although specimens in Bed C at Bovey Lane,
Devon, considered here as local variants of O. multicostata Pettitt, resemble O. boussensis. The
latter can be distinguished by its more evenly biconvex profile, more massive beak, higher
uniplication and from the Devon specimens by its less transverse outline; it somewhat resem-
bles specimens from the Plenus Subzone of Lincolnshire.

DIMENSIONS of some paratypes. Measurements in mm.

Number length width thickness Number length width thickness
BB61140 15-2 16-8 11-3 BB61144 13-3 15:5 10-9
BB61141 15-3 16-2 itile7/ BB61145 ils) 16-8 10-6
BB61142 14-1 ISP 7/ 10-7 BB61146 13-5 16:5 10-5
BB61143 13-8 16-2 10-2

Family NORELLIDAE Ager, 1959
Subfamily MONTICLARELLINAE Childs, 1959
Genus MONTICLARELLA Wisniewska, 1932

Monticlarella brevirostris (Roemer)
Pl. 18, fig. 4.

1840 Terebratula brevirostris Roemer: 41; pl. 7, fig. 7.
1852 Rhynchonella martini Mantell; Davidson: pl. 12, figs 16, 16a—d.

REMARKS. This species is thought to be of stratigraphical importance and is being investigated
with C. J. Wood. It resembles and has often been confused with Grasirhynchia martini
(Mantell), p. 91, with which it is found in the Chalk Marl at Folkestone, Dover, Eastbourne,

104 E. F. OWEN

and at Compton Bay and Rocken End on the Isle of Wight. Whether it occurs independently of
Grasirhynchia martini is not known.

It differs from other Monticlarella species in having well developed costae and in lacking the
parasulcate anterior commissure characteristic of M. carteri (PI. 5, figs 29-31) and M. jefferiesi.

MATERIAL. Numerous specimens from the Chalk Marl, rhotomagense Zone of Folkestone and
Dover, British Museum (Natural History).

Superfamily TEREBRATULACEA Gray, 1840
Family TEREBRATULIDAE Gray, 1840
Subfamily SELLITHYRIDINAE Muir-Wood, 1965
Genus SELLITHYRIS Middlemiss, 1959

Sellithyris tornacensis (d’ Archaic)
Pl. 6, figs 1-3

1841 Terebratula subundata Phillips; Roemer: 42; pl. 7m, fig. 15.

1847 Terebratula tornacensis d’Archiac: 316; pl. 18, figs 3a—e, 4a, 5a—b.

1867 Terebratula tornacensis d’Archiac; Schloenbach: 438 (38); pl. 21 (1), figs 8, 8a.
1878 Terebratula tornacensis d’Archiac; Deike: 12, fig. 3a, b.

1929 Rectithyris tornacensis (d’Archiac) Sahni: 10.

1959 Sellithyris tornacensis (d’Archiac) Middlemiss: 113.

NeoTyPE. The original material, including Terebratula tornacensis, of d’Archiac’s (1847) classic
description of the brachiopod fauna of the Tourtia has been lost (Dr Annie Dhondt, personal
communication). A neotype, from the Tourtia of Tournai, Belgium, B85066, matching the
illustrations in d’Archiac (1847) is therefore selected. Length 30-0 mm; width 28-4 mm; thickness
18-3 mm. PI. 6, figs 1-3.

OCCURRENCE. The species occurs comparatively rarely in the Essen Greensand at Kassenberg,
north Germany, in the Sable et grés du Mans at Le Mans, and in the Sables du Perche,
Broillay. The species may have been recorded as Terebratula biplicata Sowerby and Sellithyris
biplicata from the hardground sequences of Orbiquet and Goneville, Normandy in the Craie
Glauconieuse, and Juignet’s (1974) record of Sellithyris biplicata from the above localities is
correct. S. tornacensis also occurs in the greensand deposits at Aix-en-Gohelle, Artois (Parent
1893).

REMARKS. This is a distinctive species and was well described by Middlemiss (1959). Morpho-
logical variation amounts to a usually slightly more oval outline or less marked pentangulate
aspect and less acute biplication of the anterior commissure than shown in d’Archiac’s original
figure (1847: pl. 18, figs 3, 4, 5). In the Sables du Mans and the Sables du Perche, where the
species appears to be more prolific than in the Belgian type locality, and Essen Greensand,
specimens tend towards a less truncated umbo and slightly more erect beak.

PLATE 6

Figs 1-3 Sellithyris tornacensis (d’Archiac). Tourtia, Tournai, Belgium. Neotype, herein selected,
B85066, x 15.

Figs 4-6 Harmatosia crassa (d’Archiac). Tourtia, Tournai, Belgium. BB84908, x 14.

Figs 7-9 Sellithyris phaseolina (Valenciennes, in Lamarck). Cliff Section at Porte des Barques,
Charente, France. BB82286, x 14.

Figs 10-12 Rectithyris smaeiniormn sp. nov. Lower Cenomanian, mantelli Zone, White Hart sand pit,
Wilmington, Devon. Holotype BB82175, x 1.

Figs 13-15? Ornatothyris sp. Grey/Red Chalk, Speeton Cliff, N. Yorkshire. BB82346, x 15.

Figs 16-18 Concinnithyris cf. subundata (J. Sowerby). Culver Cliff, Isle of Wight. BB76462, x 15.

Figs 19-21 ‘Ornatothyris’ sp. Grey Chalk, Speeton, N. Yorkshire. BB43360, x 15.

oes a ee anglia sp. nov. Bed C, Red Rock, Hunstanton, Norfolk. Holotype

x

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 105

106 E. F. OWEN

DIMENSIONS of BM(NH) specimens. Measurements in mm.

Number and locality length width thickness
B85067, Tournai 25-0 D333} 14-8
B85068 zs 27-2 21-9 14-6
B85069 Fe 23-9 21-0 37
B85070 ‘ 24-1 18-5 11-5
B85071 $5 Dips) 25-0 16:3
B85072 . 28:3 26-6 17:3
65485, Briollay (Sarthe) 27:4 24:6 15:9
95 27-0 Daal 15-6
: ss 28:0 26:2 16:8
: 53 24-7 23S) 16-0
. 55 Daal 22:8 i139)
os 5 23-8 22:9 14:0

Sellithyris phaseolina (Valenciennes, in Lamarck)
Pl. 6, figs 7-9; Pl. 18, figs 11-13
1819 Terebratula phaseolina Valenciennes, in Lamarck: 251, No. 29.
1850 Terebratula phaseolina Valenciennes, in Lamarck; Davidson: 439; pl. 13, fig. 29.
1910 Terebratula phaseolina Val. in Lm.; Clerc & Favre: pl. 6, figs 28-33.
1971 Sellithyris biplicata Gaspard: 1—10; pl. 1, figs 1a—7c; pl. 2, figs 1-6.

LECTOTYPE. The specimen figured by Clerc & Favre (1910: pl. 6, figs 33), one of the remaining
four syntypes in the Lamarck Collection, Museum d’Histoire Naturelle, Geneva. Length
14-1 mm; width 12:4 mm; thickness 8-0 mm. Herein selected.

Like so many of the Lamarck species, there was neither clear indication of a type specimen
nor illustration of the species. Davidson (1850: 439) attempted to establish the species by
figuring a specimen from the Lamarck Collection, but this cannot be selected as lectotype
because his drawing appears to have been made up from several of Lamarck’s syntypes.

Clerc & Favre (1910: pl. 6, figs 28-33) catalogued eight specimens in the Lamarck Collection,
figuring six which were identified as A-F. Examination of these six remaining Terebratula
phaseolina specimens, in the Department of Geology, Museum d’Histoire Naturelle, Geneva,
reveals that only four can be positively identified: pl. 6, fig. 29 = specimen A, fig. 30 = specimen
C, fig. 31 = specimen D, and fig. 33 = specimen F, the lectotype.

REMARKS. At the type locality, Le Mans, large and smaller forms are found together in the
sandy, less chalky, facies; the lectotype is of the smaller variety. This does not have the marked
biplication or the elongated an inflated umbo of the larger form. Both varieties are recognized
in collections from the Marnes a O. biauriculata of Briollay (Maine et Loire) and from beds in
the Metoicoceras gourdoni Zone of Méziéres sous Ballon (Sarthe).

DIMENSIONS of BM(NH) specimens. Measurements in mm.

Number length width thickness Number length width thickness
Mayet (Sarthe): Port des Barques:

BB82280 17:3 16-1 8-8 BB82286 18-9 16:6 10-7
BB82281 18-5 16-8 10-9 BB82287 16:9 15:1 10-0
BB82282 18-2 16:4 9:7 BB82288 18-1 17:2 10-0
BB82283 17-0 14-9 9-3 BB82289 18-5 16-9 11-0
Mézieres sous Ballon:

BB82290 15:5 14:3 9-3

BB82291 13S) 12-2 6:7

Genus OVATATHYRIS nov.
TYPE SPECIES. Terebratula ovata J. Sowerby, 1812.

NaME. The name Ovatothyris appeared in a Ph.D. thesis by Dr Margaret Cox (University of
London, 1967). Her diagnosis and description have been emended and the name altered to

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 107

Ovatathyris in order to agree with the species. Credit is due to Dr Cox for the recognition of
this genus.

D1AGNosts. Outline oval, beak suberect, foramen large, circular, marginate, mesothyridid; beak
ridges well defined. Symphytium short, well exposed. Anterior commissure sulcate to slightly
paraplicate. Folding concentrated in anterior half of shell. Cardinal process low, tapering.
Crural processes high, inwardly inclined. Transverse band moderately highly arched.

EMENDED DESCRIPTION. The low, flat cardinal process, seen in transverse serial sections of the
two Ovatathyris species described here, is typical of species in the subfamily Sellithyridinae, as is
the degree of concavity of the hinge-plates. It is noteworthy that the transverse band, while
maintaining a fairly uniform breadth, is more highly arched than in Musculina, Platythyris or
Boubeithyris.

Ovatathyris remained consistently oval during growth but developed a marked sulcus
anteriorly on the dorsal valve which, in anteriorly carinate specimens, led to paraplication of
the commissure.

REMARKS. Ovatathyris differs from Boubeithyris in having an oval outline, sharper beak-ridges,
more highly developed and relatively more frequently occurring growth-lines, and shell orna-
ment of short radiating spinules. It differs internally from Boubeithyris in having slightly longer
and more acutely concave hinge-plates, and from Walkerithyris by the absence of piped hinge-
plates and in having a sulciplicate to paraplicate anterior commissure.

The genus is represented only by the two species described herein. One of these, Ovatathyris
ovata, occurs as a comparatively common fossil in the Lower Cenomanian, Mantelliceras
mantelli Zone, of the Grizzle at Wilmington, south Devon and from the Glauconitic Marl of
Warminster, Wiltshire. The other species, Ovatathyris potternensis sp. nov., also occurs in the
Glauconitic Marl at Warminster and the type locality at Potterne, Wiltshire. A variant of O.
potternensis occurs rarely in the highly fossiliferous beds within the Glauconitic Marl of the Isle
of Wight.

Ovatathyris ovata (J. Sowerby)
Fig. 15; Pl. 7, figs 19-24; Pl. 25, figs 3-4

1812 Terebratula ovata J. Sowerby: 46; pl. 15, fig. 3.

1849 Terebratula lacrymosa d’Orbigny: 99; pl. 512, figs 6-11.
1852 Terebratula ovata Sowerby; Davidson: 47; pl. 4, figs 6-13.
1874 Terebratula ovata Sowerby; Davidson: 32; pl. 2, fig. 14.

DESCRIPTION. Oval in outline with a sulcate to paraplicate anterior commissure. The large,
circular foramen truncates the massive suberect umbo, bounded by sharp beak-ridges, outlining
a short triangular interarea. A shallow dorsal sulcus originates at about half the valve length
and deepens and widens anteriorly. The pedicle valve is correspondingly carinate, a feature
commonly originating at about one-third of the valve length, producing a greater convexity
than in the brachial valve.

Shell ornament consists of step-like concentric growth-lines and short spinules which,
however, may not be preserved. When examined under a hand lens it is seen that the short
spinules are arranged along growth lines in pairs, unlike those of Arcuatothyris which are more
widely scattered. The higher magnification offered by the electron microscope (PI. 25, figs 3-4)
shows the short, paired bases of the spinules on the surface of O. ovata from the Warminster
district. Although Davidson described and illustrated these spinules on several specimens (1852:
pl. 4, figs 6-13) their preservation is not always good, probably because of abrasion, and indeed
they cannot be distinguished on the type specimen.

INTERNAL CHARACTERS. In transverse serial sections through a specimen from Warminster it can
be seen, Fig. 15, that a pedicle collar was not developed in the ventral umbo and that a low,
poorly developed cardinal process is present in section 3. The long, slender hinge-teeth first
appear on section 4 and by sections 5 and 6 are deeply inserted into the brachial valve sockets.

108 E. F. OWEN

TN am

. = ce .
26 04 S 04 Sa 0) , aess

03 03

ae _

0:2

Fig. 15 Ovatathyris ovata (J. Sowerby). Fifteen transverse serial sections through 88764 from the
Upper Greensand, Warminster, Wiltshire.

Acutely ventrally concave hinge plates, from which strong inner and outer socket ridges arose,
become progressively U-shaped anteriorly. The crural processes are long and slender, curving
gently towards one another. A highly-arched transverse band of the brachial loop occurs at
about 6:5 mm from the ventral umbo.

HOLotyPe. Sowerby Collection, B49840; from the Lower Chalk; Chute Farm, Wiltshire.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 109

DIMENSIONS of BM(NH) specimens. Measurements in mm.

Number and locality length width thickness
B492, Warminster 24-6 19-3 13-9

39 Pa 25:0 17:2 14-6

3° a 32). 20:1 18-3

a es 28-5 18-6 16-0

= 5 26:5 18-2 13-6
BB5534, south Wiltshire 32-4 25-5 18-1
B6297, Vimoutiers (Orne), France 33-8 24-4 18-1

REMARKS. The species is well known from the Warminster and Wilmington localities, but
occurs also in Bed A of the Cenomanian Limestone at Chardstock, Devon (B1522); rarely from
the Chalk Basement Bed, acutus Subzone of the rhotomagense Zone, Middle Cenomanian of
Snowdon Hill, Chard (BB82219); the Lower Chalk, Basement Bed, Osmington, Dorset
(BB82152) and from an unknown zone within the Cenomanian from Maiden Newton, Dorset,
in the J. F. Walker Collection (B25279).

In France, Ovatathyris ovata occurs rarely in the coarse sandy beds and the glauconitic marls
which crop out along the Normandy Coast from St Jouin to Cap de la Héve, where it has been
called ‘Terebratula lacrymosa d’Orbigny. Roemer’s (1840: 44) record of T. ovata from this area
is inaccurate.

O. ovata appears to be confined to the British and French Lower Chalk, but five specimens
showing a similar outline, strongly sulcate to paraplicate anterior commissure and typical
spinose ornament have been found in the Pictet Collection, Museum d’Histoire Naturelle,
Geneva from Perte du Rhone, Switzerland (CB 8, 56-58, 89). These are compared to O. ovata
and one is figured here (PI. 7, figs 19-21).

Ovatathyris potternensis sp. nov.
Fig. 16; Pl. 7, figs 10-18; Pl. 25, figs 1-2

1847 Terebratula squamosa Mantell; Davidson & Morris: 254; pl. 18, fig. 8, a, b.
1852 Terebratula squamosa Mantell; Davidson: 50; pl. 5, figs 7—9.

DIAGNOsIs. Biconvex Ovatathyris; pentagonal to oval in outline. Umbo massive, slightly pro-
duced, beak suberect, beak-ridges distinct. Interarea extensive, symphytium well exposed;
foramen large, circular, mesothyridid. Biplicate to paraplicate. Rugose ornament.

NAME. ‘From Potterne’.

DESCRIPTION. An ornament of short spinules, similar to those described for the type species,
occurs on the lateral flanks of well preserved specimens. This can be seen using a light micro-
scope but is best recorded using a scanning electron microscope (PI. 25, figs 1-2, from examples
of O. potternensis, compared with those from O. ovata, PI. 25, figs 3—4). Serial sections (Fig. 16)
compare closely with those taken through a specimen of Ovatathyris ovata from Warminster,
Wiltshire.

Ho.otyPe. BB82149; from the Upper Greensand; Rifle Butts, Potterne, Wiltshire. Length
24-3 mm; width 18-7 mm; thickness 16:0 mm. PI. 7, figs 10-12.

REMARKS.. The rugose ornament of concentric growth-lines on this species led Davidson to
believe that he was describing Mantell’s Terebratula squamosa from the Chalk Marl of Hamsey,
but a close study reveals that they lack the additional ornament seen on T. squamosa. Compare
Pl. 25, figs 1-2 (O. potternensis) with Pl. 27, figs 1-4 (Capillithyris squamosa); see also PI. 28, figs
1-4 (C. disparilis). O. potternensis also occurs in the Glauconitic Marl of Compton Bay, Isle of
Wight.

E. F. OWEN

110

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 111

/ res

os KES NURS

| QS G S S 2 : t Cy
23 03 ‘- eee. ee ‘5

0-4

03

Fig. 16 Ovatathyris potternensis sp. nov. Twenty-three transverse serial sections through 88763 from
the Upper Greensand of Potterne, Wiltshire.

PLATE 7

Figs 1-3 Boubeithyris diploplicata sp. nov. Lower Cenomanian, Vimoutiers (Orne), France. B6289,
x 1.

Figs 4-9 Boubeithyris boubei (d’Archaic). Figs 4-6, Tourtia, Tournai, Belgium. BB82370, x 1. Figs
7-9, Lower Cenomanian, Le Havre, Normandy. B10816, x 1.

Figs 10-18 Ovatathyris potternensis sp. nov. Figs 10-12, Glauconitic Marl, Potterne, Wiltshire.
Holotype BB82149, x 1. Figs 13-18, Glauconitic Marl, Compton Bay, Isle of Wight. Figs 13-15,
BB59839, x 1. Figs 16-18, BB82189, x 15.

Figs 19-21 Ovatathyris cf. ovata (J. Sowerby). ? Upper Albian, Perte du Rh6éne, Switzerland. Geneva
Museum (Nat. Hist.) coll. No. CB 57, x 14.

Figs 22-24 Ovatathyris ovata (J. Sowerby). Warminster Greensand, Warminster, Wiltshire.
BB84902, x 15.

1D) E. F. OWEN

O. potternensis can be distinguished from O. ovata by its consistently acute biconvexity and
its rugose shell ornament. It can be distinguished from other species of Terebratulidae, such as
Ornatothyris, largely by its internal characters; also the acute ventrally concave hinge-plates
and high arched transverse band of the brachial loop.

It seems likely that the species is confined to the Upper Greensand.

MATERIAL. Additional specimens from Potterne are in the Walker Collection (B25261, B25284),
the Davidson Collection (B8287—9) and the C. W. & E. V. Wright Collection (BB82148); and
88763: Fig. 16. Also from the Isle of Wight (BB59839 and BB82189).

Genus BOUBEITHYRIS Cox & Middlemiss, 1978

TYPE Species. Terebratula boubei d’Archiac, 1847.

Boubeithyris boubei (d’ Archiac)
Pl. 7, figs 4-9

1847 Terebratula boubei d’Archiac: 320; pl. 19, fig. 11.
1903 Terebratula boubei d’Archiac; Lamplugh & Walker: 252; pl. 18, fig. 5.
1978 Boubeithyris boubei (d’Archiac); Cox & Middlemiss: 419; pl. 40, figs 1-4; text-fig. 4.

Neotype. M.T.C. 10154, from the Tourtia of Tournai, Institut Royal des Sciences Naturelles de
Belgique, Brussels. Selected Cox & Middlemiss, 1978.

REMARKS. Cox & Middlemiss (1978) recognized the species in the Lower Albian of Leighton
Buzzard, Bedfordshire, and extended its geological range to Lower Albian—Cenomanian. It is
noteworthy that the ecological conditions at Leighton Buzzard during the Lower Albian were
different from any others at that time in the United Kingdom, with the possible exception of
the Melton Carstone, in Yorkshire. They were, however, very similar to those of part of the
Tournai district in Belgium.

The genus Boubeithyris is represented in the British Cenomanian by B. diploplicata sp. nov., a
species formerly referred to ‘“Terebratula biplicata’, sensu J. Sowerby (1815) and J. de C.
Sowerby (1825).

Boubeithyris diploplicata sp. nov.
Fig. 17; Pl. 7, figs 1-3; Pl. 8, figs 4-24

1815 Terebratula biplicata J. Sowerby: 201; pl. 90, figs 2-5 (non fig. 1, = Moutonithyris dutempleana
(d’Orbigny)).

1852 Terebratula biplicata Brocchi; Davidson: pl. 6, figs 34-39.

1867 Terebratula biplicata Sow.; Schloenbach: pl. 21, figs 1-3.

1871 Terebratula biplicata Sow.; Quenstedt: pl. 48, figs 61—62.

1903 Terebratula biplicata Sow.; Jukes-Browne & Hill: 123, 129.

1969 Terebratula dutemplei dOrbigny: Panow: 583; pl. CXI, figs 1-3.

D1aGnosis. Elongate-oval, to pentagonal, to subquadrate in outline. Beak short, suberect.

PLATE 8
Figs 1-3 Tropeothyris vectis sp. nov. Lower Cenomanian, Mont Ste Catharine, Rouen, France.
B6156, x 13.

Figs 4-24 Boubeithyris diploplicata sp. nov. Figs 4-6, Upper Greensand, Compton Bay, Isle of
Wight. B25281, x 13. Figs 7-9, Upper Greensand, Warminster, Wiltshire. Holotype B95597, x 14.
Figs 10-12, Lower Cenomanian, Vimoutiers (Orne), Normandy. B6289, x 14. Figs 13-15, Upper
Greensand, Rocken End, Isle of Wight. BB69002, x 13. Figs 16-18, Cenomanian, Warminster,
Wiltshire. BB76462, x 14. Figs 19-21, Glauconitic Marl, Eastbourne, Sussex. BB82349, x 14. Figs
22-24, Glauconitic Marl, Fecamp, Normandy. B35066, x 14.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE

113

114 E. F. OWEN

Foramen large, circular, mesothyridid; attrite to marginate. Anterior commissure strongly
biplicate to sulciplicate.

Cardinal process well developed, hinge-plates thin, ventrally concave, transverse band mod-
erately highly arched.

Name. ‘Doubly folded’.

DESCRIPTION. Although the species is variable it is distinguished by its very acute anterior
paraplication and strong biconvexity. The foramen is large and circular, and the beak-ridges
well defined. The ventral umbo is short and massive and the interarea poorly developed and
obscured by the dorsal umbo. Growth-lines developed throughout growth, but are particularly
evident at the shell margins.

Ho.otyre. B95597; from the Lower Cenomanian, Upper Greensand; Warminster, Wilts.
Length 28:5 mm; width 21-5 mm; thickness 20-6 mm (PI. 8, figs 7—9).

PARATYPES. B95590—602, Warminster; BB5578-89, Maiden Bradley; B6221, Undercliff, Isle of
Wight; BB6900, Rocken End, Isle of Wight; B35063, Cap de la Héve, near Le Havre, Nor-
mandy; B85265, Auberville, Normandy; B6124, Vimoutiers (Orne), France.

REMARKS. This species was often recorded as ‘Terebratula biplicata Sowerby’, but that specific
name had already been used for a Jurassic species by Brocchi (1814). Davidson (1852) followed
Sowerby in identifying the acutely biplicate species from beds of Lower Cenomanian age in the
Warminster district as T. biplicata Sowerby. It was thought that the species was confined to
localities in Devon, Dorset and Wiltshire but, because of a better understanding of its wide
variation in size and degree of convexity, it has since been recognized in the more sandy facies
of the Normandy Cenomanian beds. In Britain, the distribution has been extended to the Isle
of Wight, where it occurs in the Glauconitic Marl and from the same horizon in basal Cenoma-
nian at Cow Gap, Eastbourne. A splendid series of specimens from the Eastbourne area,
showing a wide variation in size, has been kindly lent me by Mr Simon Hutt. In this area the
species was formerly represented only by a single specimen in the C. W. Wright collection, but
now PI. 8, figs 19-21 illustrates a further example.

Other localities include Rye Hill Farm and Deverill Hill, near Warminster, and Chute Farm,
Wiltshire; a good series of specimens are in the collections of the British Geological Survey,
GSM 109902-14, 109924—30, 110036—S2 and 111015-35.

The specimens figured by Schloenbach (1867: pl. 21, figs 1-3) came from the Unterer Planer
of north Germany. The specimen figured by Panow (1968: pl. CXI, fig. la—c) came from the
Cenomanian of the Krakow district, Poland; it was part of a collection described in 1930 but
not published until revised by Biernat & Popiel-Barczyk (1969).

All the above specimens are assigned to Boubeithyris on the strength of serial sections made
from B82750 (Fig. 17). They are similar to the sections of Cox & Middlemiss (1978: 420) and of
Cox (née Pinder) (1967, unpublished Ph.D. thesis, University of London).

Subfamily RECTITHYRIDINAE Muir-Wood, 1965
Genus RECTITHYRIS Sahni, 1929
TYPE SPECIES. Terebratula depressa Valenciennes, in Lamarck, 1819.
Rectithyris wrightorum sp. nov.
Pl. 6, figs 10-12

DiaGnosis. Oval, biconvex Rectithyris. Umbo short, foramen large, circular; symphytium not
exposed. Rectimarginate.

Name. For Messrs C. W. & E. V. Wright.

DESCRIPTION. The short, massive umbo is truncated by a large circular foramen. Beak-ridges
are lacking and the interarea is poorly defined. Growth-lines are numerous but indistinct,

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 115

| |

0-4 02

Fig. 17 Boubeithyris diploplicata sp. nov. Nineteen transverse serial sections through B82750, from
the Upper Greensand of Beuzeville, near Le Havre, Normandy.

becoming clearer towards the margins. The shell is inflated in the dorsal umbo but remains
biconvex in lateral profile.

Ho.oryPe. C. W. & E. V. Wright Collection, BB82175; from the Lower Cenomanian, ‘Grizzle’;

White Hart sand pit, Wilmington, south Devon. Length 45:-3mm; width 38-9mm; thickness
18-9 mm.

116 E. F. OWEN

PARATYPE. BB82215, collected by Dr W. J. Kennedy from the same locality and horizon as the
holotype. Length 45-3 mm; width 38-9 mm; thickness 18-9mm.

REMARKS. Rectithyris wrightorum is an extremely rare species in the Lower Cenomanian of
Wilmington, but is probably the ecological equivalent of the type species R. depressa, described
from the Essen Greensand. The new species differs in its short, massive umbo, whereas the type
species, like its equivalent Terebratula nerviensis d’Archiac in the Belgian Tourtia, is character-
ized by its slender, produced umbo and well exposed symphytium. Otherwise the two species
have a similar oval outline and exhibit approximately the same degree of biconvexity.

In many ways, a closer morphological comparison is with Terebratula shenleyensis Walker,
1903, from the Lower Albian of Leighton Buzzard, Bedfordshire and redescribed with its
varieties by Dr M. Cox (1967, Ph.D. thesis), differing from T. wrightorum sp. nov. in its oval
outline and less acutely inflated brachial valve.

Internal structures remain unknown through a lack of suitable material for sectioning.

Genus TROPEOTHYRIS Smirnova, 1972
TYPE SPECIES. Tropeothyris kugusemi Smirnova.

Tropeothyris vectis sp. nov.
Fig. 18; Pl. 8, figs 1-3; Pl. 9, figs 1-6, 10-15

DIAGNOsIS. Pentagonal to broadly oval in outline. Umbo short, massive; foramen large, circu-
lar. Beak suberect, beak-ridges mesothyridid. Biplication poorly to moderately developed.

NAME. Vectis; ancient name for the Isle of Wight.

DESCRIPTION. The lateral profile is ventri-biconvex, the brachial valve being only gently curved.
A shallow sulcus starts at about mid-length of the adult dorsal valve and deepens and broadens
anteriorly; it is bordered by faint carinae. Individuals with their maximum width just anterior
to the middle of the shell tend to have a stronger biplication at the commissure. Faint growth-
lines occur between two or more well-marked growth halts at about half and two-thirds of the
adult shell length.

INTERNAL CHARACTERS. Transverse serial sections (Fig. 18) display a pedicle collar and a wide
symphytium. A low, flat cardinal process can be seen in section 7, just after the first appearance
of the dorsal umbo. Short, ventrally concave hinge-plates elongate slightly and become virgate.
Thickened crural bases give rise to long, slender, inwardly curving and rapidly diminishing
crural processes. A low arched transverse band terminates the loop approximately 8-5 mm from
the ventral umbo.

Hotorype. E. F. Owen Collection, BB82185; from the Glauconitic Marl; Gore Cliff, Isle of
Wight. Length 30-2 mm; width 22-4mm; thickness 16-6 mm.

PARATYPES. B6221 from Culver Cliff, Isle of Wight; B24944 from the Glauconitic Marl of
Folkestone Warren, Kent.

PLATE 9

Figs 1-3 Terebratula carteri Davidson (? = Tropeothyris). Lower Cenomanian, Folkestone, Kent.
Davidson Collection, B6126, x 13.

Figs 4-6, 10-15 Tropeothyris vectis sp. nov. Figs 4-6, Upper Greensand, Rocken End, Isle of Wight.
BB82331, x 14. Figs 10-12, Glauconitic Marl, Gore Cliff, Isle of Wight. Holotype BB82185, x 14.
Figs 13-15, Glauconitic Marl, Folkestone Warren. B24944, x 13.

Figs St) Tropeothyris sp. Cenomanian, Loudon Road Pit 19, near Louth, Lincolnshire. BB55022,
x iF.

Figs 7-9 Undescribed species. Lower Cenomanian, Julianka, Annopol, Poland. BB82181, x 13.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE AUT)

\ |
a SP

Fig. 18 Tropeothyris vectis sp. nov. Twenty-one transverse serial sections through BB82331, from
the Glauconitic Marl, Rocken End, Isle of Wight.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 119

BOO 6S 6:
Ol) ID

0-2 0

<4 . S P

Fig. 19 Tropeothyris sp. Seventeen transverse serial sections through a specimen from the Lower
Cenomanian of Julianka, Annopol, Poland.

REMARKS. Specimens have also been collected from Eastbourne in beds of early Cenomanian
age. Terebratula carteri Davidson (1852: 72; pl. 7, fig. 3) may belong to this genus but it differs
from the species described here in its uniplicate anterior commissure and slightly more pro-
duced umbo. An undescribed specimen which might belong with T. vectis was collected from
Pit 19, Loudon Road, Louth, Lincolnshire (PI. 9, figs 16-18). It is more regularly oval in outline
than the typical form and has a less acutely developed anterior plication. It appears to be
closely related to another undescribed species from the Lower Cenomanian of Annopol on the
Vistula, Poland (PI. 9, figs 7-9). Serial sections of a specimen from the Phosphate Mine,
Julianka, Annopol, Poland (Fig. 19) can be compared to sections through a specimen of
Tropeothyris vectis (Fig. 18) and to sections through a specimen from Loudon Road Pit,
Louth, Lincolnshire (Fig. 20). Each series has much in common, but more research is necessary
before their relationships can be elucidated.

120 E. F. OWEN

DIMENSIONS of some BM(NH) specimens. Measurements in mm.

Number length width thickness Number length width thickness

Glauconitic Marl; Rocken End, I. o. W. Mantelli Zone; Eastbourne.

C. W. & E. V. Wright Colln: W. J. Kennedy Colln:

BB69001 22:6 19:6 149 BB82303 25:99 21:9 143

BB69004 26:3 21:1 14-5 BB82304 27:3 21:4 16-1

BB69005 22:0 188 11:8 BB82111 31-2 21-4) 15-9

BB69006 246 178 13:0 BB82112 33-8 268 16-4

BB69007 Vee) IS e2 AS BB82113 31-7 25-1 15-0

BB69009 25:33 19:5 13-8 BB82114 29:0 22:8 13-6
BB82115 24-1 19-5097

Glauconitic Marl; Rocken End, I. 0. W. Glauconitic Marl; Ventnor, I. 0. W.

W. J. Kennedy Colln: F. Stinton Colln:

BB82122 27:0 20:8 16:1 BB9298 29:2 Slee 8

BB82123 DA Oe sall BB9299 26:0 20:7 12-0

BB82124 add) aesk 13) BB9300 23 Oils mls 26

BB82125 21-4 18-1 9-6 BB9301 24:4 19:1 13:5

S SO)
1-6
ey O- <a
0-4

4
5 mm a
as ie Oe

Ve Y

Nee os YY ( VAY

Fig. 20 Tropeothyris sp. Ten transverse serial sections through a specimen from the Loudon Road
Pit, Louth, Lincolnshire, of Middle Cenomanian age.

PLATE 10

Figs 1-12 Moutonithyris dutempleana (d’Orbigny). Figs 1-3, Bed B, Red Rock, Hunstanton Cliff,
Norfolk. BB82345, x 14. Figs 4-12, Cenomanian, Cambridge Greensand, Cambridge. Figs 4-6,
BB21188, x 14. Figs 7-9, B25177, x 14. Figs 10-12, variant, B25161, x 14.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 121

122 E. F. OWEN

Genus MOUTONITHYRIS Middlemiss, 1976

TYPE SPECIES. Terebratula moutoniana d’Orbigny, 1849

Moutonithyris dutempleana (d’Orbigny)
Fig. 21; Pl. 10; Pl. 11, figs 4-6; Pl. 18, figs 23-24

1815 Terebratula biplicata J. Sowerby: 201; pl. 90, fig. 1 (non figs 2-5; see p. 112).

1823 Terebratula biplicata Sow.; J. de C. Sowerby: 53; pl. 437.

1849 Terebratula Dutempliana d’Orbigny: 93; pl. 511, figs 1, 2, 3, 6, 7 (non figs 4, 5).

1855 Terebratula biplicata Brocchi; Davidson: 55; pl. 6, figs 1-30.

1960 Terebratula dutempleana d’Orb.; Pauca & Patrulius: 88; pl. 2, fig. 10.

1975 Terebratula dutempleana d@Orbigny; Middlemiss (in Dieni, Middlemiss & Owen): 186; pl. 33, fig. 2;
text-fig. 9.

1976 ‘Terebratula dutempleana d’Orbigny; Gaspard: pls 1—S.

LECTOTYPE. B61531; from the Cambridge Greensand, Castle Hill, Cambridge; figured by J.
Sowerby, 1815: pl. 90, fig. 1. Selected Middlemiss (1975).

REMARKS. Until this species was revised by Middlemiss (1975), it had been broadly interpreted,
and had become confused with interpretations of Brocchi’s Jurassic species T. biplicata Brocchi.

There is, unfortunately, some uncertainty over the exact stratigraphical horizon of the lecto-
type, since the term ‘Cambridge Greensand’ includes the Upper Albian to Middle Cenomanian.
The adopted concept of M. dutempleana (d’Orbigny), based on the lectotype, includes the
commonly-occurring terebratulid species with its greatest shell width at just over half its total
length and with a marked plication of the anterior commissure. The species is found as both
derived and indigenous fossils within the Cambridge Greensand in the Cambridge area, and as
a common fossil in the Red Rock of Hunstanton Cliff, north Norfolk. The beds within this part
of the Hunstanton section fall within the Middle to Upper Albian and have been divided into
three distinct lithological units, A, B and C, with Bed C at the bottom resting on the Carstone.
According to Dr H. G. Owen (personal communication), who has worked on ammonite species
within this series, Bed C is in the Anahoplites intermedius Subzone of the Middle Albian, Bed B
within the upper part of the cristatum Subzone, and Bed A has a lighter band representing the
Hysteroceras varicosum Subzone at the base and a darker band within the ?Callihoplites auritus
Subzone at the top. Both Beds A and B are of Upper Albian age.

The commonest variant found in Beds A and B at Hunstanton and in the Cambridge
Greensand is represented by two specimens figured here, from the Upper Albian of Hunstan-
ton (PI. 10, figs 1-3), and from the Cambridge Greensand (PI. 10, figs 4-6). Variation within the
species affects the oval outline and degree of biplication of the anterior commissure (see PI. 10,
figs 7-9 & 10-12).

Occurring with M. dutempleana in both the Cambridge and Hunstanton beds is Terebratula
obtusa J. de C. Sowerby (1825). This species was included in Davidson’s (1855) synonymy of M.
dutempleana (then described as Terebratula biplicata), but in fact the two differ considerably. T.
obtusa maintains a consistently pentangulate outline, flatter dorsal umbo and a uniplicate to
incipiently biplicate anterior commissure; its growth-lines appears to occur more frequently,
encouraging some workers to assign it to Ornatothyris. The constant association of these two
species extends beyond Britain, and can be recognized in assemblages of Terebratulidae includ-
ing forms thought to be very closely related to Moutonithyris dutempleana from beds of Ceno-
manian age at Drap, near Nice, southern France. The two species are now known to be
congeneric and it has been suggested that this association might demonstrate sexual dimor-
phism. Serial sections of specimens from the Cambridge Greensand, similar specimens from the
Upper Albian beds of the Red Rock at Hunstanton, and specimens of Moutonithyris dutem-
pleana from Hunstanton (Fig. 21) and the type locality at Cambridge all resemble each other.
Examples of Moutonithyris obtusa (Pl. 11, figs 7-9; Pl. 12, figs 4-6) from the Cambridge
Greensand can be compared with M. dutempleana (PI. 11, figs 4-6) from Bed A, Red Rock,
Hunstanton. M. dutempleana occurs uncommonly in the Upper Gault, Price’s (1877) Bed X, at
Folkestone, Kent (Pl. 18, figs 23-24).

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 123

Fig. 21 Moutonithyris dutempleana (d’Orbigny). Twenty transverse serial sections through BB82333
(Le Strange colln), from Bed A, Red Rock, Hunstanton Cliff, Norfolk.

124 E. F. OWEN

Frebold (1935), in describing a brachiopod fauna from the marine Upper Aptian of
Koldewey Island, north-east Greenland, included a specimen he called Terebratula biplicata
(1935: pl. 8, figs 1-3), the general form of which is very close to Moutonithyris dutempleana
described here from the Cambridge Greensand and Red Rock of Hunstanton. Frebold figured
an associated crushed specimen with capillate ornament (1935: pl. 8, fig. 4), which is very
similar to Capillithyris capillata (d’Archiac), the species associated with Moutonithyris species at
Hunstanton.

Cooper (1983), in his major study of terebratulacean loops, described a new genus and
species commonly occurring in the Upper Albian Red Rock of Hunstanton, Norfolk, which
had been called ‘Terebratula dutempleana d’Orbigny. This species was assigned to Mouton-
ithyris by Middlemiss (1976: 63) when he erected the genus. As a result of careful hand
dissection of the brachidia and cardinalia, Cooper was able to expose certain internal charac-
teristics in a specimen of that species which are not seen in serial sections published by Gaspard
(1976) and Middlemiss (in Dieni, Middlemiss & Owen 1975). Cooper displayed the fairly wide
concave hinge-plates attached to the ventral part of the crural bases, and the acutely high
folded and broad transverse band at the front of the brachial loop. The description of these
characters conflicts with that of Moutonithyris given by Middlemiss (1976) in which he stressed
the low arching of the transverse band and the cuneate, keeled hinge-plates. However, trans-
verse serial sections of Mountonithyris dutempleana (d’Orbigny) and M. anglia sp. nov. (p. 126)
from the Red Rock of Hunstanton (Figs 21, 22) do not support Cooper’s description of the
ventral attachment of the outer hinge-plates but do agree in the presence of a short, broad loop,
slightly expanded anteriorly and with an acute transverse band, characters shown also in
transverse serial sections by Gaspard (1976) of another specimen from the Red Rock of
Hunstanton.

Cooper described as new Biplicatoria feruginea, the type species of his new genus, and B.
hunstantonensis from the top bed (Bed 3 = Bed A) of the Upper Albian, Red Rock. I contend
that Moutonithyris dutempleana (d’Orbigny) occurs predominantly in Bed 3 (Bed A) and that
M. anglia sp. nov. is restricted to Bed 1 (Bed C) at the base of the Red Rock.

Thus while acknowledging Dr Cooper’s work I do not accept that M. dutempleana should be
transferred to Biplicatoria, nor do I believe that the specimens figured by him (Cooper 1983: pl.
20, figs 21-27; pl. 21, figs 1-6) as Biplicatoria hunstantonensis and B. feruginea are other than
variants of Moutonithyris dutempleana (d’Orbigny), although this requires confirmation.

MATERIAL. Numerous specimens of Moutonithyris dutempleana (d’Orbigny) are housed in the
British Museum (Natural History), British Geological Survey, and the Sedgwick Museum,
Cambridge. Specimens of M. obtusa also occur in the same institutions but are less numerous.
Six specimens from the Upper Albian, Red Rock of South Cave, Humberside are in the
collections of the British Museum (Natural History), nos 68962-7, and show a very close
resemblance to the type specimen from the Cambridge Greensand.

DIMENSIONS. The following measurements (in mm) are from BM(NH) specimens from Beds A
and B in the Red Rock, Hunstanton, Norfolk, collected by Mr H. Le Strange.

PLATE 11

Figs 1-3 Ornatothyris sabinensis (Parent). Tourtia, Tournai, Belgium. BB82188, x 13.

Figs 4-6 Moutonithyris dutempleana (d’Orbigny). Bed A, Red Rock, Eieneentont Cliff, Norfolk.
BB82180, x 14.

Figs 7-9 Moutonithyris obtusa (J. de C. Sowerby). Cambridge Greensand, Cambridge. BB6006, x 1.

Figs 10-12 Concinnithyris davidsoni (Rollier). Tourtia, St Aybert, Belgium. Int. Sci. roy. Belg. I.G.
8261, x 15.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 125

126 E. F. OWEN

Number length width thickness Number length width thickness
BB82350 B72 308 21-6 BB82355 34-0 26:3 19-1
BB82351 34-0 PTS) 20-2 BB82356 31-1 DPS) 18-9
BB82352 34-5 25-0 19-1 BB82357 30-2 23-6 18-0
BB82353 30:8 Pil 19-9 BB82358 27-9 23-4 17-1
BB82354 31-1 ISPS) 19-0 BB82359 29-3 23-4 17-2

Moutonithyris anglia sp. nov.
Fig. 22; Pl. 6, figs 22-24

D1AGNosis. Consistently oval in outline. Maximum width at half valve length. Strongly ventri-
biconvex. Uniplicate to incipiently biplicate. Umbo truncated by large circular foramen; beak
suberect.

NAME. ‘England’.

DESCRIPTION. Because of their similarity, this species has, until now, been compared with M.
dutempleana. Moutonithyris anglia, however, has a consistent oval outline and a very marked

pnb e |
\ Ne

Fig. 22 Moutonithyris anglia sp. nov. Twenty-one transverse sections through B2221 from Bed C,
Red Chalk, Hunstanton Cliff, Norfolk.

5mm

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 7

even biconvexity. The anterior commissure is uniplicate to faintly biplicate, never showing the
acute biplication of M. dutempleana. The umbo is massive and dominated by the large circular
foramen with permesothyridid beak-ridges. M. anglia does not have a labiate foramen like that
of M. dutempleana, and the symphytium is never exposed.

INTERNAL CHARACTERS. Transverse serial sections (Fig. 22) show that this species, like all species
assigned here to Moutonithyris, has the same low, poorly developed cardinal process, and the
short, ventrally concave initial hinge-plates which lengthen, becoming almost horizontal and
keeled before being deflected dorsally. The terminal ends of the hinge-plates thicken, giving rise
to V-shaped bases which change anteriorly into the descending branches of the brachial loop.
This elongates ventrally, giving off inwardly curving crural processes, which diminish anter-
iorly. The transverse band is a low-arched, comparatively broad W-shaped ribbon. This struc-
ture is quite commonly associated with a feature called the ‘sub-loop skirt’ by Cox &
Middlemiss (1978) and probably resulted from coalescence of spicules within the mantle in the
living specimen. Oddly, this character has been observed in specimens of M. dutempleana and
M. obtusa from Beds A and B and in specimens from the Cambridge Greensand, but does not
seem to occur in sectioned specimens from Bed C described here as M. anglia.

Ho.ortyPe. Collected by H. Le Strange, BB82262; from Bed C of the Red Rock, Hunstanton
cliffs, Norfolk. Length 40-1 mm; width 30-2 mm; thickness 24:2 mm. PI. 6, figs 22—24.

PARATYPES. Ten specimens from the type locality and horizon, BB82263-72. Dimensions given
below.

REMARKS. This is the only Moutonithyris species occurring in Bed C of the Red Rock. It can be
distinguished from other species of Moutonithyris by its consistent elongate-oval outline with
its maximum width at half the valve length, large circular foramen, and gently uniplicate to
faintly biplicate anterior commissure.

DIMENSIONS of paratypes. Measurements in mm.

Number length width thickness Number length width thickness
BB82263 29-3 20-9 19-1 BB82268 31-2 24-5 16:5
BB82264 36-6 27-6 20-4 BB82269 32-4 25-6 17-4
BB82265 33-1 26:1 20-3 BB82270 30-8 Dal 18-2
BB82266 30-2 23-4 18-4 BB82271 27-9 23-0 17-2
BB82267 31-1 22-4 V7eil BB82272 29-9 23-2 15-9

Subfamily NERTHEBROCHINAE Cooper, 1983
Genus HARMATOSIA Cooper, 1983

Harmatosia crassa (@ Archiac)
Pl. 6, figs 4-6

1847 Terebratula tornacensis var. crassa d’Archiac: 318; pl. 18, figs 8a—d.
1983 Harmatosia crassa (d’Archiac) Cooper: 196; pl. 19, figs 13-22; pl. 29, figs 8-11; pl. 67, figs 5, 10, 11.

REMARKS. This species was originally described as a variety of Terebratula tornacensis by
d’Archiac (1847) from the Tourtia of the Tournai district, Belgium. Cooper (1983) chose it as
type species of his new genus Harmatosia and an example, from the type locality, is illustrated
here (PI. 6, figs 4-6), for comparison with Sellithyris tornacensis (d’Archiac) (p. 104; Pl. 6,
figs 1—3).

RANGE. Cenomanian.

128 E. F. OWEN

Family GIBBITHYRIDIDAE Muir-Wood, 1965
Subfamily GIBBITHYRIDINAE Muir-Wood, 1965
Genus CONCINNITHYRIS Sahni, 1929
TYPE SPECIES. Terebratula obesa J. de C. Sowerby 1823.

EMENDED DIAGNOSIS. Elongate-oval to subcircular in outline. Umbo massive, beak slightly
incurved to erect, truncated; foramen small to large, circular, permesothyridid, rarely labiate;
beak ridges indistinct. Anterior commissure rectimarginate, to uniplicate, to biplicate.

Cardinal process low, flat, poorly developed. Hinge-plates fairly long, ventrally convex from
anterior ends of which crural bases develop. Crural processes high, tapering, inwardly curving.
Thin descending branches of brachial loop with low arched transverse band, sometimes with
small depression or sulcus posteriorly.

RANGE. Upper Albian to Cenomanian.

REMARKS. In his description of the brachial loop of Concinnithyris, Cooper (1983: 183) gave the
stratigraphical range as Neocomian to Turonian and included the species C. albensis
(Leymerie), C. obesa (Sowerby), C. burhamensis Sahni and C. rouenensis Cooper, 1983. Although
disagreeing with Cooper’s (1983) wider stratigraphical range, I believe the genus may be rep-
resented within the basal beds of the Lower Turonian in France.

The interpretation of this genus depends upon the interpretation of the type species, T. obesa
J. de C. Sowerby. As the holotype is the only representative of this species in the Sowerby
Collection, and is from an unrecorded horizon at Norton Bavant, Wiltshire, Sahni (1929) must
have based his description of the genus on this and other specimens from different localities
which he identified as T. obesa. Although he states the range of the type species to be Turonian,
he dissected a specimen from Eggardon Hill, Dorset, B24914, which is probably also T. obesa.
This locality has been reinterpreted by Kennedy (1970) as of jukesbrownei assemblage age, at
the top of the Acanthoceras rhotomagense Zone of the Middle Cenomian. Sahni’s age determi-
nation was probably based upon data recorded by d’Orbigny (1847) who cited conspecific
specimens from the “Turonien Superieur’ of Rouen, and it is indeed a comparatively common
fossil within the A. rhotomagense Zone in the Craie de Rouen.

Concinnithyris obesa (J. de C. Sowerby)
Fig. 23; Pl. 12, figs 7-12; Pl. 14

1823 Terebratula obesa J. de C. Sowerby: 54; pl. 438, fig. 1.
1850 Terebratula obesa J. de C. Sowerby; d’Orbigny: 105; pl. 513, figs 1-3 (non fig. 4).
1929 Concinnithyris obesa (J. de C. Sowerby) Sahni: 12; pl. 1, figs 1-9; pl. 8, figs 1—2.

Ho.otyPe. Sowerby Collection, B49832; from the Lower Chalk; Norton Bavant, Wiltshire.
Length 48-9 mm; width 36-0 mm; thickness 34-9 mm.

PLATE 12
Figs 1-3 Ornatothyris latissima Sahni. Plenus Subzone, South Ferriby, S. Humberside. BB82202,
x 15.

Figs 4-6 Moutonithyris obtusa (J. Sowerby). Cambridge Greensand, Cambridge. IGS unregistered
specimen, x 15.

Figs 7-12 Concinnithyris obesa (J. Sowerby). Figs 7-9, Cenomanian, Storridge Hill, Chardstock,
Devon. BB84905, x 15. Figs 10-12, Bed C, Cenomanian, Shapwick Farm, near Uplyme, Dorset.
BB82172, x 14.

Figs 13-15 Ornatothyris cf. pentagonalis Sahni. First Inoceramus Bed, Lower Cenomanian, Hun-
stanton Cliff, Norfolk. BB82302, x 14.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 129

5mm

Fig. 23 Concinnithyris obesa (J. de C. Sowerby). Seventeen transverse serial sections through
BB82332, from the Lower Chalk, 1 m above the Cenomanian Basement Bed, White Nothe, Dorset.

REMARKS. Sahni (1929: 12) gave an adequate description of the species based on the Sowerby
type and accompanied this with two dissected specimens (1929: pl. 8, figs 1-2), one of which,
B49884, he called a plesiotype from “The Chalk’ of Lewes, Sussex. This specimen, from an
unspecified horizon within the Chalk, should only be assigned to Concinnithyris obesa (J. de C.
Sowerby) with doubt.

Sahni did not recognize variability in the species and so considered only large, almost perfect,
but miserably few, specimens matching the holotype as representative of C. obesa. In fact,
thorough investigation of specimens collected from the Lower Chalk of Dorset, Wiltshire and
the Isle of Wight shows that the range of variation includes specimens which do not match the
holotype in their general dimensions; these may be steeper-flanked and with subparallel out-
lines, may be anteriorly constricted and also less acutely biconvex. A range of variants rep-
resenting the species is figured here (PI. 14, figs 1-15), including the holotype (figs 1—3).

Specimens from a bed less than one metre above the phosphatic conglomerate comprising
the Cenomanian Basement Beds are always highly calcareous, non-phosphatized, and whitish
in colour. This bed occurs extensively at Snowdon Hill, Chard, Somerset; White Nothe, Ring-
stead and Durdle Cove, Dorset; Charstock, Devon; and in the Isle of Wight. From amongst

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 131

1:8

02

Fig. 24 Concinnithyris davidsoni (Rollier). Seventeen serial sections through B5070, from the Upper
Greensand of Warminster, Wiltshire. One section also shown at greater enlargement.

such specimens a variant is recognized by its flatter valves, large foramen, and more acute
biplication of the anterior commissure.

All variants, including specimens like the type, have been recognized in collections from the
Craie de Rouen, France, from where the species was collected and described by d’Orbigny
(1850) as from his ‘Etage Turonien’. Since the work of Kennedy & Hancock (1970), Juignet &
Kennedy (1976) and others, it is now known that the Craie de Rouen is of Middle Cenomanian
age.

Davidson (1852: 53; pl. 5, figs 13-15) interpreted C. obesa as the biplicate terebratulid which
occurs rarely in the Warminster Greensand. Apart from their large size the two forms are
dissimilar and it seems strange that Davidson should have made this decision, especially since
Sowerby’s original specimen of T. obesa was available to him. Because of this misinterpretation
Rollier (1911: 28) changed the name of the Warminster species to Terebratula davidsoni, and

132 E. F. OWEN

this species occurs also in the Tourtia of Tournai, Belgium (Pl. 11, figs 10-12). It has been
confused with Terebratula sabinensis Parent, 1893 (Pl. 11, figs 1-3), which should probably be
placed in Ornatothyris. T. sabinensis differs from C. davidsoni in its evenly oval outline, suberect
and truncated umbo and lack of biplication of the anterior commissure.

Transverse serial sections through a young specimen of Concinnithyris davidsoni (Fig. 24)
display the comparatively short ventrally deflected inner hinge-plates, the long incurved crural
processes and the low W-shaped transverse band of the brachial loop, and should be compared
with the series for the type species (Fig. 23).

DIMENSIONS. Measurements in mm.

Locality number length width thickness
Isle of Wight BB69017 28-9 23-9 19-3
BB82080 44-8 33-3 34-9
Ringstead BB69013 B21 24-8 21-0
Osmington BB45120 45:2 35-8 33-1
Durdle Cove BB69088 30-4 23-1 20:5
BB69085 30:7 26:0 19-6
Snowdon Hill BB82364 40-6 34-9 26:1
BB82365 31-8 29-2 20:5
BB82366 29-4 23-9 18-7
Rouen, France BB82367 44-9 38-6 34-4

Concinnithyris subundata (J. Sowerby)
Figs 25-27; Pl. 6, figs 16-18; Pl. 15, figs 4-18; Pl. 16

1813 Terebratula subundata J. Sowerby: 47; pl. 15, fig. 7.

1929 Concinnithyris subundata (J. Sowerby) Sahni: 17; pl. 1, figs 10-16 (non fig. 17); pl. 8, figs 11-12.
1929 Concinnithyris subundata (J. Sowerby) Sahni: 17; pl. 1, figs 10-16 (non fig. 17); pl. 8, figs 11-12.
1929 Concinnithyris burhamensis Sahni: 16; pl. 2, figs 8-13; pl. 8, figs 22-23.

EMENDED DESCRIPTION. The beak is suberect epithyridid with a modest circular foramen.
Symphytium not exposed. The outline of the shell varies from subpentagonal (PI. 15, fig. 4) to
subcircular (Pl. 15, fig. 13), to elongate-oval (Pl. 16, fig. 1). An even greater degree of biconvexity
is common for adults, but diminishes in some variants. Rarely a shallow dorsal median sulcus
occurs, usually bordered by faint carinae, forming a weakly biplicate anterior commissure,
which otherwise is gently uniplicate.

INTERNAL CHARACTERS. Apart from the early stages in the development of the hinge-plates, and
their shorter length than those of the type-species, other internal features closely resemble those
of C. obesa; compare Fig. 23 with Fig. 25.

LECTOTYPE. Sowerby Collection, B49827, figured by J. Sowerby 1813: pl. 15, fig. 7 (Pl. 16, figs
1—3). Herein selected.

Sowerby’s (1813: 47) original description appears to have been based on several specimens,
but illustrated by only one (1813: pl. 15, fig. 7), a strongly biconvex, elongate specimen having,
according to Sowerby, ‘a fleshy red colour’. Today there are six specimens named C. subundata
in the Sowerby Collection (B49826—31). When Sahni (1929) included this species in his new
genus Concinnithyris he selected as what he called a “Neanic Holotype’ specimen B49826
(Pl. 15, figs 1-3), but mentioned no other Sowerby specimen as any part of a type series. This
specimen closely resembles neither the above original Sowerby figure nor most of Sahni’s own
illustration of the species (1929: pl. 1, figs 10-15). In addition the matrix is glauconitic, and it is
thus probably from the Chalk Rock rather than from the ‘softish Chalk at Warminster’
(Sowerby 1813), which is probably Lower Chalk in that area.

Specimen B49827 resembles Sowerby’s fig. 7 more closely than does B49826. It also fulfills
most of Sahni’s own criteria for his description of C. subundata, as well as matching his
illustrations (1929: pl. 1, figs 10-15). The species is well known from the Lower Chalk, from
which specimen B49827 probably came, and for these reasons I select it as Lectotype of C.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 133

03 03 0-2

5mm

Fig. 25 Concinnithyris subundata (J. Sowerby). Twenty-one transverse serial sections through
BB82335, from the Lower Chalk, Southerham Gray Pit near Lewes, E. Sussex.

134 E. F. OWEN

subundata (J. Sowerby). The specimen is not now flesh coloured, but such colouration tends to
fade from newly collected specimens.

REMARKS. Measurements of specimens from the Basement Beds in Dorset and Wiltshire, and
some from French localities, reveal that in many there is considerable variation in the size of
the foramen. In addition these specimens tend to be less biplicate or have a simple uniplicate
anterior commissure. As far as can be detected from the comparatively few that have been
serially sectioned, these differences are not reflected internally.

Dimensions of BM(NH) specimens. Measurements in mm.

Number length width thickness Number length width thickness
Burham, Kent: Punfield, Dorset:
B9449 33-4 26:1 22:1 BB69104 30-8 26:2 18-2
B9450 34-4 26-2 22-1
: Evershot, Dorset:

Pees ae BB82163 325 280) 209

BB82164 29-8 25:5 19-2
Folkestone, Kent: BB82165 32-7 28:7 22:8
B22876 30-4 26:3 19-2 BB82166 30-5 24-1 18-4
BB5602 30:5 25:4 19-5 :

; : : Burwell, Cambridge:

BB5603 34-1 27-2 20:5 B24968 32-1 6-7 16-8
Oxted, Surrey: es Die 25:0 16:1
B58233 37-6 31:3 22:9 re 26:7 24:5 16-9
B44820 30-1 27-1 18-2 ee
Bes822 2 2 IZ B35074 260 27mm.
Chinnor, Oxon: 5 26:3 23-1 15:6
B47274 31:5 26:5 19-0 BB82167 28-6 25:3 18-0
Guildford, Surrey: St Jouin, Normandy:
B10634 29:3 25:4 15-4 BB82168 29-8 25-8 18-9
Culver, Isle of Wight: BBS2169 — 22:7 14-5
BB68996 39-4 33-8 22:0
BB81153 35:7 29-0 20-3

C. subundata also occurs in the Grey Chalk and the Grey/Red Chalk of Speeton, Yorkshire
and examples from the BM(NH) collections are BB43357—-59, BB45141—49, BB69113-17,
BB69119, BB69147—59 and BB82774-809.

Genus ORNATOTHYRIS Sahni, 1929
Type species. Terebratula sulcifera Morris, in Davidson & Morris 1847.

REMARKS. In his original description of the genus, Sahni (1929) gave the geological range as
Cenomanian—Senonian. Within the nine species he described he distinguished two morphologi-
cal groups by differences in rugose shell ornamentation. One group, which he termed ‘toti-
rugose’, included all the Cenomanian species in which both valves are covered by prominent

a —————————o

PLATE 13

Figs 7-9 Ornatothyris sulcifera (Morris). Cenomanian, Fulbourn, Cambridge. BB6017, x 13.

Figs 1-3, 10-12 Ornatothyris cf. sulcifera (Morris). Figs 1—3, Cenomanian, Cassis, southern France.
BB82174, x 14. Figs 10-12, Middle Cenomanian, Grove Mill Pit, Hitchin, Herts. BB82190, x 13.
Figs 4-6, 13-18 Ornatothyris sp. Figs 4-6, Craie de Rouen, Céte Ste Catharine, Rouen. BB82200,
x 14. Figs 13-18, Brunni Alp, Bernese Oberland, Switzerland. Geneva Nat. Hist. Mus. Figs 13-15,

C.B. 366, x 14. Figs 16-18, C.B. 369, x 1.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 135

136 E. F. OWEN

WIDTH

*= Lectotype

LENGTH

Fig. 26 Length x width graph and regression line for 102 specimens of Concinnithyris subundata
(J. Sowerby) from England.

THICKNESS

LENGTH

Fig. 27 Length x thickness graph and regression line for 102 specimens of Concinnithyris subundata
(J. Sowerby) from England.

PLATE 14

Figs 1-15 Concinnithyris obesa (J. de C. Sowerby). Figs 1-3, Lower Chalk, Norton Bavant, Wilt-
shire. Holotype B49832, x 1. Figs 4-6, Craie de Rouen, Céte Ste Catharine, Rouen. BB84903, X1.
Figs 7-9, Middle Cenomanian, Ringstead, Dorset. BB45120, x 1. Figs 10-12, Craie de Rouen,
Rouen. B5101, x 1. Figs 13-15, Middle Cenomanian, Isle of Wight. BB84904, x 1.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 137

138 E. F. OWEN

growth-lines. His second group, which included all the species from the Micraster coranguinum
Zone of Greenhithe, Kent, was termed ‘partirugose’ because this ornamentation was present
only anteriorly.

Five of the six totirugose species came from the Cambridge district; the sixth was said to
have come from an unknown locality but a second specimen was figured from Warminster,
B49828, a locality from which the genus has never since been recorded. Although somewhat
vague, the localities quoted by Sahni from the Cambridge area are reasonably recognizable, but
it is not known for certain from exactly which horizon each of Sahni’s holotypes originates. The
Lower Chalk of the Cambridge district covers many of the zones within the Cenomanian, the
lowest part being represented by the Cambridge Greensand, considered to be of carcitanense
equivalent age, i.e. lowest Cenomanian (Carter & Hart 1977: 75). The highest level ranges to
the Plenus Marl.

It is likely that O. sulcifera is a variable species and that most of Sahni’s other species from
the same Cambridge area are merely variants. The possible exceptions are O. pentagonalis and
O. latissima, which appear as consistent morphological forms; in particular O. pentagonalis
(Pl. 12, figs 13-15) is found in the “Paradoxica’ Bed of the “White Chalk’ of Hunstanton Cliff, i.e.
low in the succession. O. latissima (Pl. 12, figs 1-3), on the other hand, appears to be the
dominant species of an assemblage containing remanié Orbirhynchia multicostata and Monti-
clarella jefferiesi in pockets of green clay in the conglomeratic erosion surface below the Black
Band at Elsham and Caistor in Lincolnshire, and at Hillington and Heacham in Norfolk. This
is high in the succession, in the top part of Bed IX of Bower & Farmery (1910) in the upper
Holaster trecensis Zone.

Ornatothyris differs from Concinnithyris externally by having a clearly developed ornament of
rugose concentric growth-lines. Internally it differs in its less developed cardinal process,
shorter and thicker hinge-plates, and dorsally pendant and shorter brachial loop. It also differs
in this way from Moutonithyris, but Terebratula obtusa J. de C. Sowerby (1825) from the
Cambridge Greensand, and previously called Ornatothyris obtusa, is here assigned to
Moutonithyris: see p. 122.

Ornatothyris sulcifera (Morris)
Fig. 28; Pl. 13, figs 1-3, 7-12
1847 Terebratula sulcifera Morris, in Davidson & Morris: 354; pl. 18, fig. 7.

1855 Terebratula sulcifera Morris; Davidson: 64; pl. 7, figs 17—20.
1929 Ornatothyris sulcifera (Morris) Sahni: 48; pl. 7, figs 1-10; pl. 10, figs 25-27.

Ho.ortypPe. Davidson Collection, 50822; from the Lower Chalk; near Cambridge.

REMARKS. Sahni figured four specimens from the Lower Chalk of Fulbourn (1929: 48) to
illustrate the species. Another Fulbourn, Cambridge specimen, said to have been collected from
the subglobosus Zone by L. Barrett, is 36-2 mm long, 28-0 mm wide and 23-2 mm thick (BB6017;
Pl. 13, figs 7-9).

The species appears to be confined to eastern England and Cassis, southern France, where
specimens occur in Lower Cenomanian ironstained limestone deposits. One of these (BB82147)
is figured (Pl. 13, figs 1-3); it is 30-4mm long, 26:-7mm wide and 18-9mm thick. It is broader

PLATE 15

Figs 1-15 Concinnithyris subundata (J. Sowerby). Figs 1-3, specimen suggested by Sahni (1929) as
lectotype of this species, but not here accepted. ?Chalk Rock (see p. 132). Sowerby colln, B49826,
x 1}. Figs 4-6, Middle Cenomanian, Oxted, Surrey. B44830, x 14. Figs 7-9, large example from
Lower Chalk, Burham, Kent. 96789, x 14. Figs 10-12, Chalk Basement Bed, Snowdon Hill,
Chard, Somerset. BB82171, x 14. Figs 13-15, Craie de Rouen, Céte Ste Catharine, Rouen, France.
BB82167, x 15.

Figs 16-18 Concinnithyris cf. subundata (J. Sowerby). Cenomanian, Grey/Red Chalk, Speeton Cliff,
N. Yorkshire. BB69046, x 14.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 139

140 E. F. OWEN

O04 O6

Eos
Ke yo OR < . /)
eo 05

O03 o4 0:3 Ly)
\ | \ (1) \ ae
ee Ca 3 eee

—

Fig. 28 Ornatothyris sulcifera (Morris). Seventeen transverse serial sections through BB82338 from
the Lower Chalk, Fulbourn, Cambridge.

PLATE 16

Figs 1-12 Concinnithyris subundata (J. Sowerby). Figs 1-3, lectotype, here selected. 7Lower Chalk
(see p. 132). Sowerby colln, B49827, x 15. Figs 4-6, Eggardon Grit, Eggardon Hill, Dorset.
BB82170, x 14. Figs 7-9, Middle Cenomanian, Glynde, E. Sussex (C. burhamensis aspect).
BB82347, x 14. Figs 10-12, Grey/Red Chalk, Speeton Cliff, N. Yorkshire. BB84906, x 14.

Figs 13-15 Concinnithyris cf. subundata (J. Sowerby). Pit 3, Hallington, Louth, Lincolnshire.
B55145, x 14.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 141

142 E. F. OWEN

and more circular in outline and has more numerous, evenly spaced growth-lines than the
holotype; the umbo is also more massive than in typical O. sulcifera or any other species within
the genus. The circular foramen is perhaps slightly proportionately larger with a marked
pedicle collar.

A British specimen collected by C. J. Wood, from the upper third of the subglobosus Zone of
Grove Mill Pit, Cadge Lane, Hitchin, Hertfordshire, is similar to the Cassis specimens in
outline, convexity and type of concentric ornament, but differs in having a less massive umbo
which is truncated by a large circular foramen. This specimen is 25-0mm long, 223mm wide
and 15-7 mm thick.

INTERNAL CHARACTERS. Seventeen transverse sections (Fig. 28) through an unregistered speci-
men from the Lower Chalk of Fulbourn, Cambridge, display a well developed pedicle collar
guarding a large foramen in the ventral umbo. Short, thickened horizontal hinge plates flatten
proximally and curve gently towards the floor of the dorsal valve, developing short pendant
plates which thicken anteriorly, turning into long, slender, inwardly curving crural processes.
The comparatively short brachial loop widens very slightly, terminating in a broad, high-
arched transverse band.

The hinge-plates have very strong inner but weaker outer socket-ridges. The hinge-teeth fit
into shallow, rounded sockets.

Subfamily CAPILLITHYRIDINAE Cooper, 1983
Genus CAPILLITHYRIS Katz, 1974

Capillithyris squamosa (Mantell)
Pl. 18, figs 14-16; Pl. 27

1822 Terebratula squamosa Mantell: 132.

1847 Terebratula squamosa Mantell; Davidson & Morris: 254; pl. 18, figs 8, 8a, 8b.
1847 Terebratula disparilis d’Orbigny (pars): pl. 512, fig. 17 only.

1852 Terebratula squamosa Mantell; Davidson (pars): 50; pl. 5, figs 5, 6 only.

1874 Terebratula squamosa Mantell; Davidson: 33; pl. 2, fig. 5.

1978 Platythyris squamosa (Mantell) Middlemiss: 39, figs 11, 12, 16 (9).

Hovotype. Mantell Collection, B8292; from the Chalk Marl of Hamsey, near Lewes, Sussex.

REMARKS. Middlemiss (1978) fully described this species in his paper on the genus Platythyris so
no further description is necessary here.

There has been some confusion between this species and Arcuatothyris arcuata (Roemer),
p. 144, since both occur in the more highly calcareous beds in southwestern England, particu-
larly in the phosphatic basement beds of the B group on the coast of south Devon. This
confusion is made worse by the presence of a third species, which is also somewhat homo-

PLATE 17

Figs 1-3 Gemmarcula carantonensis (d’Orbigny). Upper Cenomanian, Port des Barques, Charente,
France. BB45960, x 2.

Figs 4-6 Gemmarcula menardi (Lamarck). Sable du Mans, Le Mans (Sarthe). BB35152, x 2.

Figs 7-9, 28-30 Arenaciarcula beaumonti (d’Archaic). Figs 7-9, Cenomanian, Drap, near Nice.
BB85424, x 14. Figs 28-30, Tourtia, Tournai, Belgium. BB45970, x 2.

Figs 10-15 Dereta pectita (J. Sowerby). Figs 10-12, ?7Middle Cenomanian, near Warminster, Wilt-
shire. B25263, x 2. Figs 13-15, Upper Greensand, from type locality at Horningsham, Wiltshire.
BB4199, x 14.

Figs 16-18 Dereta pectita ballardensis subsp. nov. Upper Albian, S. dispar Zone, Ballard Point,
Swanage, Dorset. Holotype BB82098, x 2.

Figs 19-24 Gemmarcula canaliculata (Roemer). Figs 19-21, Tourtia, Tournai, Belgium. BB82191,
x 14. Figs 22-24, Lower Cenomanian, White Hart sand pit, Wilmington, Devon. BB82193, x 2.

Figs 25-27 Dereta incerta (Davidson). Glauconitic Marl, Chardstock, Devon. BB84907, x 2.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 143

144 E. F. OWEN

eomorphic: it was described as a subspecies Platythyris squamosa disparilis (d’Orbigny) by
Middlemiss (1978), but I consider it a true species C. disparilis. C. squamosa can be distin-
guished from C. disparilis by its flatter valves, wider hinge-line and less massive umbo which is
truncated by a comparatively large circular foramen. The outline of C. squamosa is oval,
although many individuals display an anterior tapering of the valves.

Both species have a marked ornament of growth-lines which are evenly spaced and show
unusual cuspidate leading edges (Pls 27, 28) and impersistent fine longitudinal striae: in the
larger C. disparilis, this ornament tends to become more zigzag towards the margins of the
shell (PI. 28, figs 3, 4). This characteristic ornament distinguishes Capillithyris from Arcua-
tothyris, and is particularly useful since C. squamosa resembles Arcuatothyris arcuata (Roemer)
in outline. The former has flatter valves and lacks an anterior sulcation of the brachial valve, a
feature of Roemer’s original illustration (Roemer 1841: pl. 8, figs 18a—c).

Family BOREIOTHYRIDAE Dagys, 1968
Genus ARCUATOTHYRIS Popiel-Barczyk, 1972

Arcuatothyris arcuata (Roemer)
Pl. 26

1841 Terebratula arcuata Roemer: 44; pl. 8, figs 18a—c.

1847 Terebratula rugulosa Morris, in Davidson & Morris: 153; pl. 18, figs 5a—c.
1847 Terebratula disparilis d’Orbigny: 100; pl. 512, figs 12, 13, non figs 16, 17.
1852 Terebratula rugulosa Morris; Davidson: 49; pl. 4, figs 14, 14a.

1866a Terebratula arcuata Roemer; Schloenbach: 451; pl. 21, fig. 12.

1878 Terebratula arcuata Roemer; Deike: 15, figs 6a—d.

1969 Terebratella arcuata (Roemer) Panow: 594; pl. 110, figs 8, 9.

1970 Terebratula rugulosa Morris; Kennedy: 634, 660.

1974 Arcuatothyris arcuata (Roemer) Katz: 268; pl. 87, figs 18a—d, 20a—c.
1978 Terebratula rugulosa Morris; Owen: 148, 151.

1978 Terebratula rugulosa Morris; Middlemiss: fig. 16, 3a—d.

TYPE SPECIMEN. Roemer’s original type specimen is missing but it seems likely from his draw-
ings (Roemer 1841: pl. 8, figs 18a—c) that the species is a senior synonym of T. rugulosa Morris,
1847. If a neotype is to be considered it should come from the conglomeratic horizon known as
the Essen Greensand.

As the presumed junior synonym Terebratula rugulosa Morris, the species is known in
Britain from the Glauconitic Marl of Chardstock, Devon, the holotype being in the Davidson
Collection, B6130; it was refigured by Davidson (1852: pl. 4, fig. 14) and Middlemiss (1978: fig.
15/2).

REMARKS. The species can be distinguished from Capillithyris squamosa (Mantell) by its steeper
flanks, and shallow sulcus anteriorly on the brachial valve. It differs from Ovatathyris ovata in

PLATE 18

Figs 1-3 Capillithyris capillata (d’Archaic). Tourtia, Tournai, Belgium. B46351, x 13.

Fig. 4 Monticlarella brevirostris (Roemer). Chalk Marl, Folkestone. 82144, x 5.

Figs 5-10 Concinnithyris sp.? Two similar specimens. Figs 5-7, Bed B, Red Rock, Hunstanton,
Norfolk. BB82145, x 2. Figs 8-10, Lower Cenomanian, Brockterbek, north Germany. BB82146,
x 2.

Figs 11-13 Sellithyris cf. phaseolina (Valenciennes, in Lamarck), possibly an elongated variant.
Upper Cenomanian, Charente. BB82212, x 13.

Figs 14-16 Capillithyris squamosa (Mantell). Lower Chalk, St Jouin, Normandy. BB85611, x 13.

Figs 17-22. Modestella geinitzi (Schloenbach). Figs 17-19, Lower Chalk, Folkestone. BB82214, x 2.
Figs 20-22, Essen Greensand, Essen, Germany. BB82213, x 2.

Figs 23-24 Moutonithyris dutempleana (d’Orbigny). Upper Gault, Price’s Bed X, Folkestone.
Fig. 23, B1132, x 15. Fig. 24, B2231, x 15.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 145

146 E. F. OWEN

PLATE 19

Figs 14 Immature Terebratulina spp. Cambridge Greensand, Cambridge. Figs 1—2, thought to be
T. etheridgei nom. nov. (= T. triangularis Etheridge, non Tate). B7021, SEM x 40. Figs 3-4,
thought to be T. nodulosa Etheridge. BB82353 and 82354 respectively, SEM x 35.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 147

its general outline and lack of ventral carina, poorly defined beak-ridges, smaller foramen and
less marked brachial sulcus. It differs from both these species in its characteristic shell ornament
of spinules which are arranged roughly radially over the surface of the valves (PI. 26, figs 1—4)
but with areas between some growth-lines devoid of spinules. These are thought to represent
times when growth at the mantle edge was too rapid for the development of spinules. This
ornament differs from that of Ovatathyris in its distribution over the surface of the shell. In A.
arcuata the spinules are more strongly developed and interdigitate, whereas in Ovatathyris they
do not normally overlap with those from adjacent bands.

This species occurs with Capillithyris squamosa and C. disparilis in the phosphatic chalky
conglomerates of the Basement Beds of the Lower Chalk at Evershott, Eggardon Hill, Toller
Porcorum, Toller Fratrum and Beaminster, Dorset; and at Bovey Lane, Wilmington sand pit
and from the coastal sections at Humble Point and the Pinnacles, Devon, where it occurs in
Beds B and C of the Cenomanian.

MATERIAL. Abundant material from the above localities is in the Kennedy Collection, BM(NH).
Additional material includes: BB76507 from the Glauconitic Marl, Chardstock, Devon;
BB82221—5 from the Lower Chalk Basement Bed, Snowdon Hill, Chard, Somerset; BB82361
from Horn Hill Quarry, Beaminster; BB82362—5 from the White Hart sand pit, Wilmington;
BB82260 from Hutchin’s Pit, Wilmington, Devon; and B6131 from the Craie de Rouen, Céte
Ste Catharine, Rouen, France.

Family CANCELLOTHYRIDIDAE Thomson, 1927
Subfamily CANCELLOTHYRIDINAE Thomson, 1927
Genus TEREBRATULINA @Orbigny, 1847

Terebratulina etheridgei nom. nov.
IAL, Sy ites, 17/3 IAL, WG) saya 1h 2

Nom. Nov. pro Terebratulina triangularis Etheridge, 1881: 148; pl. 13, fig. 15; non Terebratulina
triangularis Tate, 1880: 160; pl. 8, fig. 7.

Hovorype. Jukes-Browne Collection, BGS no. 3156; from the Cambridge Greensand.

REMARKS. Unfortunately, Etheridge’s original species name Terebratulina triangularis had pre-
viously been used by R. Tate to describe a species from the Lower Tertiary of South Australia,
so although long used to describe specimens from the Cambridge Greensand, it must now be
changed.

The species can be distinguished from other Terebratulina species by its consistently triangu-
lar outline. This is very largely on account of the slightly more extended hinge-line which adds
greatly to the posterior width of the shell and increases the interarea. It can be further distin-
guished from T. imbricata sp. nov. (p. 148) in having non-lamellose growth lines. The radial
ribbing, while exhibiting similar bifurcation and intercalation to that seen on T. imbricata sp.
nov. and T. nodulosa Etheridge, is less prominent.

MATERIAL. Numerous specimens from the J. F. Walker Collection and many other sources in
the British Museum (Natural History), British Geological Survey and the Sedgwick Museum,
Cambridge.

Terebratulina nodulosa Etheridge
Pl. 5, fig. 16; Pl. 19, figs 3, 4; Pl. 21, figs 1-4.

1881 Terebratulina nodulosa Etheridge: 148; fig. 13.
1903 Terebratulina nodulosa Etheridge: Jukes-Browne & Hill: 204, 215, 224.

EMENDED DESCRIPTION. Circular to oval in outline with strong radiating ribs (14-18) orna-

148 E. F. OWEN

mented by well-developed nodules. Short umbo truncated by large circular foramen. Average
size about 9-5 mm in length, 8-6 mm wide and 2-4mm thick.

NEoTYPE. B.G.S. specimen No. 117521; from the Totternhoe Stone, Burwell Rock; Burwell,
Cambridgeshire (PI. 21, figs 2-4). Herein selected.

REMARKS. Etheridge (1881) described two species of Terebratulina from the Cambridge area,
one of which, T. triangularis (now T. etheridgei) has been discussed on p. 147; the other, T.
nodulosa, was described from the Totternhoe Stone of Cherry Hinton. Unfortunately, the type
specimen, which was represented in Etheridge’s illustration (1881: fig. 13) is lost. A specimen
subsequently thought to have been the holotype was found to be T. triangularis from the
Cambridge Greensand; a neotype is therefore selected.

OccURRENCE. The nodular ribs of T. nodulosa are distinctive amongst species of Terebratulina.
Specimens are known from the Upper Albian at Hunstanton and from the Lower Chalk of
Folkestone and Dover, and eight from the Lower Chalk, Bed V, of Lydden Spout, Folkestone
(B25004). One in the collections of the B.G.S. (no. Hr.9030) came from the top of the O.
mantelliana Band of Kennedy [ = Tottenhoe Stone Horizon] from the Dover Pier borehole, at
a depth of 202 ft. In addition Terebratulina nodulosa appears in the faunal list given by Jukes-
Browne & Hill (1903: 40) from Bed 4 (Bed V of Price) in the Lower Chalk at Folkestone.

Terebratulina imbricata sp. nov.
Pl. 22

Terebratulina ornata (Roemer); auctt.
Terebratulina nodulosa Etheridge; auctt.

DIAGNOosIs. Oval to subcircular in outline. Umbo short, foramen small, circular. Ornament of
about 18 coarse ribs, but up to 20 on some specimens. Growth lines imbricate. Rectimarginate.
Average length 6-7 mm, width 5-8 mm, thickness 2:2 mm.

Name. ‘Overlapping like tiles’.

DESCRIPTION. This small Terebratulina is characterized by the frequent marginal ventral bifur-
cation and dorsal intercalation of its ribs, and by its lamellose growth-lines which produce an
imbricate surface ornament. Additional ornament occurs as faint rounded papillae lying prin-
cipally between the costellae (Pl. 22, figs 2-4). The rib nodules, so characteristic of many species
of Terebratulina, occur faintly where weak growth lines cross the ribs, especially during early
stages of growth and around the umbonal regions. No folding or sulcation of the valves
occurred at any stage of growth, and the anterior commissure remains rectimarginate.

Ho.otype. Collection of C. W. & E. V. Wright, BB82196; Lower Pink Band (Bower &
Farmery Bed V equivalent), Lower Chalk; Rifle Butts Pit, Market Weighton, Humberside (PI.
22, figs 1-4).

PARATYPES. Three other specimens from the same collection, BB82197—9; and five specimens
from the same horizon at Clapham’s Limeworks, near Louth, Lincolnshire, in the collections of
the B.G.S., Nos 99058-62.

PLATE 20

Figs 1-2 Terebratulina protostriatula sp. nov. Chalk Marl, Hamsey, E. Sussex. BB82416. Fig. 1, shell
surface ornament on brachial valve. SEM x 30. Fig. 2, nodulose ornament from umbonal region.
SEM x 40.

Figs 3-4 Terebratulina striatula (Mantell). Fig. 3, shell surface ornament of brachial valve of holo-
type now assigned to the Upper Chalk. B457, SEM x 35. See also PI. 5, figs 7-9. Fig. 4, shell
surface of the brachial valve of specimen from the Upper Chalk, cortestudinarium Zone, Wiltshire.
The angular nature of the striae contrasts with those of T. protostriatula from the Lower Chalk.
BB82417,SEM x 35.

149

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE

150 E. F. OWEN

Specimens have also been collected by K. Evans from the Lower Pink Band at Hubbard’s
Vale, near Louth, Lincolnshire.

REMARKS. As Bower & Farmery’s collection was destroyed during World War II, the identifica-
tion of the species they (Bower & Farmery 1910) called ‘Terebratulina ornata’ in the Lower
Pink Band remains unknown. Terebratulina nodulosa Etheridge, originally from the Totternhoe
Stone, Cambridge, has more recently been collected from beds above the Totternhoe Stone, and
Jukes-Browne & Hill (1903: 224) recorded it from the Totternhoe Stone and younger beds in
Lincolnshire. T. nodulosa is thus the species most likely to have been the one called T. ornata
Roemer by Bower Farmery, because their nodulose shell ornament is so similar; I believe that
T. imbricata sp. nov. is a distinct species, which may prove stratigraphically important.

Terebratulina protostriatula sp. nov.
Pl. 5, figs 1-6, 13-15; Pl. 20, figs 1-2

Terebratulina striatula (Mantell); auctt.
1826 Terebratulina striatula J. de C. Sowerby: 69; pl. 536, figs 3, 4 (non Mantell).
1852 Terebratulina striata, sensu Wahlenberg; Davidson: pl. 2, figs 25, 27, 28.

D1aGnosis. Elongate-oval, uniplicate, evenly biconvex Terebratulina. Symphytium well exposed,
foramen large, circular. Ornament of rounded bifurcating costellae.

NAME. ‘Forerunner of striatula’.

DESCRIPTION. Frequent growth-lines interrupt the longitudinal costellae. In young and some
well preserved adults the ornament is nodulose in the umbonal region of each valve. The umbo,
though slightly produced, is more massive or less constricted than in species from the Middle
and Upper Chalk.

Variation occurs in the degree of slight anterior sulcation, a tendency towards a more
spatulate general outline, and in the density of costellation.

Hovorype. Mantell Collection, B6038; from the Chalk of Hamsey, Sussex.

PARATYPES. One from the Lower Cenomanian of Wilmington, Devon, BB82261; 12 from the
Chalk Marl, Dover, Kent, B29707; 1 from the Lower Chalk, Eastbourne, Sussex, B4804; 1 from
Bluebell Hill, Burham, Kent, B9458; 2 from Peter’s Quarry, Burham, Kent, B44805-—6; 2 from
the Glauconitic Marl of Warminster, Wiltshire, B54862-3; 5 from the Glauconitic Marl, St
Lawrence, Isle of Wight, B75; 1 from the Lower Chalk of Newington’s Chalk Pit, SE of Glynde
railway station, BB862; 4 from the Lower Chalk of Balcombe Pit, Glynde, Sussex, BB43940-3 ;
5 from the lodge to Newtimber Place, Newtimber Church, Sussex, BB43965—9; 1 from the
limeworks at Chinnor, Oxfordshire, B47302; 5 from the Upper Greensand of Deverill Hill,
Dorset, B7003; 1 from the Upper Greensand, Ventnor, Isle of Wight, B10676; 6 from the Chalk
Marl, Hamsey, Sussex, BB6038—43; 3 from the chalk above the Basement Bed, Punfield Cove,
Swanage, BB76500-2; 15 from the Grey Chalk, Bed V (Price), Lydden Spout, Folkestone, Kent,
B25003; 2 small individuals from the Cambridge Greensand, B7002; 3 from the Glauconitic
Marl, Rocken End, Isle of Wight, BB76451—3; 1 from Bed 6 of Jukes-Browne (Lower Chalk),
from a cutting in railway between Dover and Folkestone, BB82363; 8 from the Lower Chalk,
Le Havre, France, B35181; 2 from Cap Blanc Nez, France, B700; 3 perfect examples from Cran
d’Escalles, Cap Blanc Nez, Boulonnais, France, BB82364-6.

PLATE 21

Figs 1-4 Terebratulina nodulosa Etheridge. Fig. 1, Middle Cenomanian, Floteberg, Germany.
BB82350, SEM x 12. Figs 2-4, Totternhoe Stone, Burwell, Cambridge. Neotype here selected,
BGS coll. no. 117521. Fig. 2, SEM x 12. Figs 3-4, enlargements of shell surface to show nodular
ornament. Fig. 3, SEM x 25. Fig. 4, SEM x 50.

Figs 5-6 Terebratulina triangularis Etheridge. Cambridge Greensand, Cambridge. BB82413. Fig. 5,
SEM x 10. Fig. 6, enlargement of shell surface ornament, SEM x 30.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 151

152 E. F. OWEN

REMARKS. This is the species which Mantell (1822) had in mind when he described a specimen
from the Chalk Marl of Hamsey in Sussex. Unfortunately he figured (1822: pl. 25, figs 7, 8, 12) a
specimen which, to judge from the adhering matrix and general morphological features, orig-
inated from the Upper Chalk. As no other specimens were involved at the time of description,
Mantell’s specimen (PI. 5, figs 7-9; Pl. 20, fig. 3) must be regarded as the holotype of Terebratu-
lina striatula (B457); it is 21-4mm long, 17-4mm wide, and 9:3 mm thick. It is larger than and
differs in outline from any specimen I have examined from the Lower Chalk. In addition it has
a marked ventral median sulcus extending from the umbonal region and becoming slightly
deeper and wider anteriorly. The surface ornament of fine costellae is like that found on
specimens of Terebratulina from the Micraster cortestudinarium Zone of the Coniacian from
downland areas and other chalk exposures in both southeastern and southwestern England.
Photomicrographs of T. striatula (P|. 20, figs 3, 4) and T. protostriatula sp. nov. (PI. 20, figs 1, 2)
show clearly that the Upper Chalk species has longitudinal costellae which are transversely
quadrate in section, while those of the Lower Chalk species are rounded. Mantell’s specimen
presumably originated from the Upper Chalk of Hamsey, and his name should therefore in
future be restricted to specimens from the Upper Chalk.

The name Terebratulina striatula cannot therefore be applied to another Terebratulina
species from the Lower Chalk. The present form is a species of Terebratulina occurring in the
Coniacian of various localities in England, often quoted erroneously as T. striata (Wahlenberg).
Wahlenberg’s species, however, was originally described from the Upper Maastrichtian of
Sweden; the original description is brief (Wahlenberg 1821: 61) and there is no illustration
available. Specimens in the British Museum (Natural History), obtained by exchange from
Sweden, are very large, 40-50 mm in length, and quite distinct from any species of Terebratu-
lina from the British Upper Chalk.

Davidson (1852) used Terebratulina striatula in the broader interpretation of Sowerby (1829:
69), which included both Cretaceous and Tertiary forms, and applied it to a specimen collected
from the London Clay of Sheppey, despite his contention that many Mesozoic and Tertiary
species should be assigned to T. striata Wahlenberg. Elliott (1938) included T. striatula in the
synonymy of his species Terebratulina wardenensis.

T. protostriatula sp. nov. differs from T. auriculata Roemer (1841), from the Hilsconglomerat
of the Essen district in north Germany. Roemer’s illustration (1841: pl. 7, fig. 9) shows a small,
very flat uniplicate specimen with a slightly extended hinge line and a large circular foramen.
The growth-lines are not very marked and the costellae appear somewhat coarser than those of
any British Cenomanian specimens I have examined. Roemer’s name has not been used for a
specimen of Terebratulina since d’Orbigny (1847) used it in his classic work on the Cretaceous;
furthermore, the exact horizon within the Hilsconglomerat of Essen is not known, the beds
ranging from Neocomian to Cenomanian in age.

For these reasons I now erect T. protostriatula, rather than using Roemer’s name auriculata,
for the species formerly known under the erroneous name Terebratulina striatula Mantell and
commonly found in the Lower Chalk of Britain and the European continent.

OTHER MATERIAL. Numerous specimens from the Lower Chalk of Britain and the Continent are
in the British Museum (Natural History) and the British Geological Survey.

PLATE 22

Figs 1-4 Terebratulina imbricata sp. nov. Lower Pink Band, Rifle Butts Pit, Market Weighton,
Humberside. Holotype, BB82196. Fig. 1, shell ornament of imbricate growth-lines and additional
intercostal papillae. SEM x 15. Fig. 2, intercalation of costae and bifurcation. SEM x 35. Fig. 3,
imbricate growth-lines at margin. SEM x 38. Fig. 4, small intercostal papillae. SEM x 45.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 153

154 E. F. OWEN

Superfamily ZEILLERIACEA Allan, 1940
Family ZEILLERIIDAE Allan, 1940
Genus MODESTELLA Owen, in Casey 1961

Modestella geinitzi (Schloenbach)
Pl. 18, figs 17-22

1866a Magas geintzi Schloenbach: 575.

1866b Magas geinitzi Schloenbach; Schloenbach: pl. 2, figs 4-8.
1903 Magas geinitzi Schloenbach; Jukes-Browne & Hill: 67, 91.
1972 Modestella sp.; Popiel-Barczyk: 134; pl. 3, figs 1-3, 7a—c.

DESCRIPTION. Small zeilleriid brachiopod approximately 8mm long, 8mm wide and 6mm
thick. Maximum width at about shell mid-length. Pentagonal in outline with short umbo, sharp
suberect beak, distinct beak-ridges and relatively large circular foramen. Conjunct deltidial
plates exposed. The concave interarea is not extensive. Anterior commissure sulcate to sulco-
carinate. The dorsal sulcus starts halfway down the valve and deepens slightly anteriorly. Two
or three well developed step-like growth-lines occur anteriorly at about two-thirds the length of
the shell.

Ho.otyPe. The whereabouts of Schloenbach’s original specimen is unknown.

REMARKS. Schloenbach (1866a: 575) described, without figuring, the species as Magas geinitzi
from the Cenomanian “Griinsand von Quedlinburg’ of north Germany, but later the same year
(1866b: pl. 2, fig. 4-8) he figured an example from the same locality.

British examples have been quoted by British authors as Magas geinitzi Schloenbach,
although differing from the German specimens in their slightly larger overall dimensions, the
German specimens being approximately 6mm long, 6mm wide and about 4mm in thickness.

Popiel-Barczyk (1972: 134) referred Polish examples of this species to Modestella sp., illus-
trating a series of transverse serial sections of a specimen from the Lower Cenomanian of
Annopol, Poland. The geological range of the genus was thus extended from Lower Albian,
from which it was first described, to Lower Cenomanian. No transverse serial sections were
given with the original description (Owen, in Casey 1961) but were provided later (Owen 1963:
199-203) in a more comprehensive description of the genus. Popiel-Barezyk’s (1972) serial
sections show an even more complete series of the internal characters of this species.

Jukes-Browne & Hill (1903: 257) recorded the species as Magas geinitzi ? Schloenbach from
the A. mantelli Zone of northwestern France and as Magas geinitzi Schloenbach from the
Lower Chalk of Norfolk, Isle of Wight and west Sussex. Oddly enough, they do not quote it
from Folkestone, where it occurs abundantly, nor does the species appear among the brachio-
pods listed from either the Lower or Middle Cenomanian of Folkestone by Kennedy (1969:
532-535). This is probably because some homoeomorphy exists between this species and Capil-
lithyris squamosa (Mantell) (p. 142), which also occurs commonly at Folkestone.

MATERIAL. Numerous specimens in the British Museum (Natural History), including the fol-
lowing from the Chalk Marl of Hamsey, Sussex, BB3436—43; from the Lower Chalk of
Folkestone, B25016, BB82214; from a similar horizon at Rougefort, Liques, Pas de Calais,
B35200; and from the Greensand of Essen, north Germany, BB82213.

The species appears to be confined to localities in Britain and northern France with a marly

PLATE 23

Figs 1-4 ‘Argyrotheca’ megatrema (J. de C. Sowerby). Cambridge Greensand, Cambridge; species
formerly assigned to Argiope by authors. Shell surface ornament resembles that of an immature
Terebratulina sp. and supports the additional ornament of intercostal papillae seen in T. imbricata
(pl. 22, fig. 4) from the Lower Pink Band of Yorkshire. BB6097, SEM x 22, x 16, x 50, x 100
respectively.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE

156 E. F. OWEN

limestone facies. The original occurrence in the Greensand of Quedlinburg and the specimens
from the Annopol district of Poland may be exceptions to this preference, although little is
known about the sediments from either of these two localities.

Family MEGATHYRIDIDAE Dall, 1870
Genus ARG YROTHECA Dall, 1900

‘Argyrotheca’ megatrema (J. de C. Sowerby)
Piss hee 27: Pl23 ele24

1836 Terebratula megatrema J. de C. Sowerby: 343; pl. 18, fig. 3.
1852 Argiope megatrema (J. de C. Sowerby) Davidson: 102; pl. 12, figs 31, 32, 34.
1852 Argiope buchii de Hagenow; Davidson: 102; pl. 12, fig. 33.

EMENDED DESCRIPTION. Small, biconvex, transversely oval terebratellid with short, almost
straight, massive ventral umbo truncated by a large circular foramen. Well defined beak-ridges
border a fairly extensive triangular ventral interarea. A disjunct delthyrium is poorly developed.
Shell ornament consists of 16-18 strong radiating costae with few intercalations. Shell surface
covered with raised papillae or nodules. Fine endopuncta are more readily observed on worn
parts of the exterior and internally.

Anterior commissure rectimarginate to incipiently sulcate, but without evidence of a corre-
sponding brachial fold.

INTERNAL STRUCTURES. Pedicle valve. There are neither supporting dental lamellae for the short
massive hinge-teeth, nor is there any indication of a median septum.

Brachial valve. There is no differentiated cardinal process, but a myophore exists between the
inner socket ridges. The broad fused hinge-plates, with massive rounded inner and outer socket
ridges, form a shallow hinge-trough supported by a low, comparatively short, median septum
extending anteriorly to support the wide descending branches of the brachial loop at about the
mid-length of the valve.

TYPE SPECIMEN. J. de C. Sowerby (1836) originally described Terebratula megatrema from the
Upper Greensand of Warminster, figuring a specimen (pl. 18, fig. 3) which is not in the Sowerby
Collection at the British Museum (Natural History) because, like so many of the brachiopods
figured in the same publication, it never formed part of his personal collection. Thus at present
no type specimen is recognized.

REMARKS. Although never regarded as a common species from the Upper Greensand and
Lower Chalk, “Argyrotheca megatrema occurs more commonly in collections from the Cam-
bridge Greensand. Fifty-five specimens collected by J. F. Walker from the Cambridge district
can be added to the three in the Davidson Collection at the British Museum (Natural History);
one (38073), from the Lower Chalk of Kent, was figured by Davidson (1852: pl. 12, fig. 33),
having been given to him by S. P. Woodward.

On the same plate, Davidson (1852: pl. 12, fig. 37) reproduced von Hagenow’s (1842) original
figure of Argiope bronni from the Upper Chalk of north Germany, which he considered a
synonym of T. buchii von Hagenow (1842). Both of these are regarded as generically and
specifically distinct from A. megatrema (J. de C. Sowerby). Argiope bronni was cited by Popiel-
Barczyk & Smirnova (1978) as type species for their new genus Bronnothyris from the Maas-
trichtian of north Germany and Poland, distinguished from Argyrotheca and all other

PLATE 24

Figs 1-3 ‘Argyrotheca’ megatrema (J. de C. Sowerby). Cambridge Greensand, Cambridge. B5247.
Fig. 1, internal structure showing broad but broken brachial loop, large hinge-teeth and extensive
hinge-line. SEM x 15. Fig. 2, enlargement to show broken supporting septal pillar and hinge-
trough with endopuncta. SEM x 34. Fig. 3, further enlarged to show shallow but broad hinge-
trough, thickened inner socket-ridges and endopuncta. SEM x 65.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE S37)

158 E. F. OWEN

megathyrid genera by the presence of rudimentary or incipient ascending branches of the
brachial loop. It can be further distinguished from ‘Argyrotheca’ megatrema (J. de C. Sowerby)
by its extended hinge-line, fewer and non-bifurcating costae, more triangular outline and the
absence of papillae on the shell exterior.

The presence or absence of these papillae in the Megathyrididae is thought by Nekvasilova
(1983) to be the result of some secondary condition, such as mode of preservation. Her explana-
tion is that the papillae are calcite infillings of large puncta protruding above the shell surface
and forming a papillate pseudosculpture. But why should this condition affect species within
the Megathyrididae and leave other species of punctate genera unaffected? It is more likely that
this is part of the primary ornamentation.

Family DALLINIDAE Beecher, 1893
Subfamily Uncertain
Genus ARENACIARCULA Elliott, 1959

Arenaciarcula beaumonti (d’ Archiac)
Pl. 17, figs 7-9, 28-30

1847 Terebratula Beaumonti d’Archiac: 331; pl. 21, figs 12-14.

1867 Terebratula (?) Beaumonti d’Archiac; Schloenbach: 461; pl. 22, figs 3—S.

1869 Trigonosemus kiprijanovi Hofman: 25; pl. 5, figs 18-21.

1871 Terebratula orbicularis Sow.; Quenstedt: 293; pl. 45, figs 63-69.

1874 Terebratella Beaumonti (d’Archiac) Zareczny: 177; pl. 2, fig. 3.

1916 Terebratella Beaumonti (d’Archiac); Ravn: 22; pl. 4, fig. 8.

1969 Terebratella beaumonti (d’Archiac); Panow: 593; pl. 112, fig. 5.

1972 Oblongarcula beaumonti (d’Archiac) Popiel-Barczyk: 127; pl. 1, figs 1—5; pl. 4, fig. 3.
1977 Arenaciarcula beaumonti (d’Archiac) Owen: 236; pl. 2, figs 7a—c.

REMARKS. This species was reviewed by Owen (1977) and the geographical distribution was
given as Britain, Belgium, Poland, Germany, Denmark and the U.S.S.R., to which can now be
added southern France. The specimen figured (PI. 17, figs 7-9), from the Cenomanian near
Nice, is an especially large variant, being larger than any of the specimens collected from
Belgium or Britain, but closely resembling the specimen figured by Panow (1969) from the
Krakow district of Poland.

Family TEREBRATELLIDAE King, 1850
Subfamily TRIGONOSEMINAE Elliott, 1965
Genus DERETA Elliott, 1959
TYPE SPECIES. Terebratella pectita (J. Sowerby, 1816).
Dereta pectita (J. Sowerby)

Figs 29-30; Pl. 17, figs 10-15

1816 Terebratula pectita J. Sowerby: 83; pl. 138, fig. 1.

1819 Terebratula pectita Sow.; Lamarck: 255.

1822 Terebratella pectita (Sow.) Brongniart, in Brongniart & Cuvier: pl. 9, fig. 3.
1838 Terebratula pectita Sow.; von Buch: 168; pl. 16, fig. 12.

PLATE 25

Figs 1-2 Ovatathyris potternensis sp. nov. Glauconitic Marl, Potterne, Wiltshire. BB82149. Fig. 1,
shell surface. SEM x 25. Fig. 2, shell margin showing immature remnants of spinules. SEM x 25.
Figs 3-4 Ovatathyris ovata (J. Sowerby). Cenomanian, Warminster, Wiltshire. Fig. 3, shell surface
showing arrangement of spinules along margin. BB6012, SEM x 30. Fig. 4, surface of similar

specimen showing spinules on brachial valve arranged in pairs. BB6009, SEM x 25.

159

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE

160 E. F. OWEN

1847 Terebratella pectita (Sow.); dOrbigny: 120; pl. 517, figs 16—20.
1852 Terebratella pectita (Sow.); Davidson: 26; pl. 3, figs 29-33.
1871 Terebratula pectita Sow.; Quenstedt: 267; pl. 44, figs 104—S.
1959 Dereta pectita (J. Sowerby) Elliott: 147.

1965 Dereta pectita (J. Sowerby); Elliott: H853, fig. 736, 2a—c.

1977 Dereta pectita (J. Sowerby); Owen: 246; pl. 1, figs 8a—c.

Ho.otyPe. Sowerby Collection, B61622; from the Upper Greensand of Horningsham, Wilt-
shire. Figured J. Sowerby (1816: pl. 138, fig. 1). Length 19-9mm; width 20-0mm; thickness
14-0mm.

20°

WIDTH

* = Holotype

LENGTH

Fig. 29 Length x width graph and regression line for 65 specimens of Dereta pectita (J. Sowerby)
from SW England.

REMARKS. This species was redescribed, with additional data from transverse serial sections, by
Owen (1977); he mentioned (1977: 248) a variant from the Upper Albian, dispar Zone of
Punfield Cove near Swanage, Dorset. It is redescribed below as Dereta pectita ballardensis
subsp. nov.

D. pectita pectita (J. Sowerby) can be distinguished from the new subspecies by its evenly
subcircular outline, equal biconvexity of valves, marked but regularly spaced growth-lines and
comparatively fine costation. It has a more clearly defined triangular interarea and a slightly
wider hinge-line.

PLATE 26

Figs 1-4 Arcuatothyris arcuata (Roemer). Chalk Marl, Folkestone, Kent. BB82352. Fig. 1, shell
surface ornament of spinules showing indiscriminate arrangement and direction. SEM x 10. Fig. 2,
area of accelerated growth between two growth-lines devoid of mature spinules. SEM x 25. Fig. 3,
shell surface ornament showing interdigitation of spinules and relative length. SEM x 30. Fig. 4,
ornament and growing edge of shell to show shortening of spinules. SEM x 50.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 161

162 E. F. OWEN

THICKNESS

5 10 15 20 30

LENGTH

Fig. 30 Length x thickness graph and regression line for 65 specimens of Dereta pectita (J. Sowerby)
from SW England.

Dereta pectita ballardensis subsp. nov.
Pl. 17, figs 16-18

Name. ‘From Ballard Point’.

DESCRIPTION. Subquadrate in outline, this subspecies is distinguished by its coarse costae and
very prominent growth-lines. The umbo is massive and the beak erect to slightly incurved. The
symphytium is not exposed. The beak-ridges are obscured by the heavy ornament. The ventral
interarea is short and poorly defined. The anterior commissure is rectimarginate to incipiently
ligate, a faint sulcus in the ventral valve being opposed by a similarly faint sulcus in the anterior
part of the dorsal valve.

HoLortyPe. BB82098; from the Upper Albian, dispar Zone of Ballard Point, near Punfield Cove,
Swanage Bay, Dorset (PI. 17, figs 16-18).

OTHER MATERIAL. Specimens in the British Museum (Natural History): from Lower Cenoma-
nian, Melcombe Bingham, B25288; from Horningsham, B61622-3; and from the Warminster
district BB4199, B25264. From the Upper Albian of Punfield Cove, Dorset, BB82099-105
(George Bate Colln).

Specimens in the Sedgwick Museum, Cambridge: from Punfield Cove, B7304; from Duns-
combe, Devon, B7291—3; from Pinhay Cliff, B7502-8.

A unique specimen in the J. F. Walker Collection, B25289, was found in the Upper Green-
sand (basal bed) in an old orchard at Chelborough, Evershot, Dorset. It is somewhat worn but
probably belongs to D. pectita ballardensis.

PLATE 27

Figs 1-4 Capillithyris squamosa (Mantell). Chalk Marl, Folkestone, Kent. BB82351. Figs 1, 2,
surface showing ‘capillate’ ornament. SEM x 15, x 25 respectively. Figs 3, 4, views of a growth
edge to show intercapillary cuspidation of the ornament of concentric growth-lines. SEM x 200,
x 420 respectively.

163

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE

E. F. OWEN

164

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 165

Dereta incerta (Davidson)
Pl. 17, figs 25-27

1852 Trigonosemus incerta Davidson: 31; pl. 4, figs Sa—c.
1871 Terebratella incerta (Davidson) Quenstedt: 268; pl. 44, fig. 106.

EMENDED DESCRIPTION. Small species of Dereta. Average dimensions: length 8-4mm; width
8-0mm and thickness 5-4mm. Hinge-line straight, ventral interarea broad, triangular; symphy-
tium well exposed. Foramen circular, comparatively large. 28 to 30 rounded costellae with
intercalations and marginal bifurcation. One to two well-marked growth-lines.

Ho.otyPe. Davidson Collection, BB82145; from the Glauconitic Marl, Chardstock, Devon.
Figd Davidson 1852: pl. 4, fig. Sa—c.

REMARKS. The only published records are those of Davidson (1852) and Quenstedt (1871), but
specimens have been found in Sussex by C. T. Gaster and in Dorset by W. J. Kennedy. In
addition three possible specimens were collected from Bed B, White Cliff, Devon, and two from
the same horizon at Little Beach, Beer Head, Devon by T. Grimsdale, B53190, B53191, B53192
respectively. These appear to be somewhat broader than the type specimen and could be dwarf
forms of D. pectita. Two specimens slightly larger than the type specimen and with coarser
costae were collected by C. W. & E. V. Wright from Downlands Beach, Devon (BB59325) and
the Glauconitic Marl of Compton Bay, Isle of Wight (BB59317).

In the Meyer Collection in the Sedgwick Museum, Cambridge there are five specimens, two
from Dunscombe from Meyer’s Beds 10-12, B7308—9, and three from Bed 13 at White Nothe,
Dorset, B20677-9.

The species is assigned here to Dereta entirely in consideration of its external morphology, as
nothing is known about the internal structures. Some of the specimens recorded here as
belonging to this species may, after further work, require reassignment.

Records of Dereta incerta show that most specimens were collected from a more highly
calcareous or chalky lithology than that in which D. pectita is found, which may account for its
wider distribution.

Superfamily THECIDEACEA Gray, 1840
Family THECIDEIDAE Gray, 1840
Subfamily THECIDEINAE Gray, 1840
Genus THECIDIOPSIS Oehlert, 1887

Thecidiopsis essensis (Roemer)
Pl. 5, figs 18-19

1837 Thecidea digitata Sowerby; Bronn: 664; pl. 30, fig. 4.

1840 Thecidea essensis Roemer: 36 (cit. Bronn 1837: pl. 30, fig. 4).

1854 Thecidea digitata (= T. essensis Roem.); Roemer: 136.

1959 Thecidiopsis (Thecidiopsis) essensis (Roemer) Backhaus: SO; pl. 4, fig. 8-9; pl. 5, fig. 1.

RemaRKS. In his original description of Thecidea essensis, Roemer (1840: 36) cited the illustra-
tion by Bronn (1837: pl. 30, fig. 4) which, although poor by present-day standards, is sufficiently
distinctive to allow differentiation of this species from other forms within the Cretaceous. The
shell is transversely oval in outline. The brachial valve is approximately 10-12mm long and
between 12 and 14mm wide. From the small collection of single valves and other material, it is
possible to estimate that complete adult pedicle valves would be about 16mm long and wide.

PLATE 28

Figs 1-4 Capillithyris disparilis (d’Orbigny). Cenomanian, Rouen, France. B35081. Figs 1, 2,
cuspidation of capillate ornament similar to that seen in C. squamosa. SEM x 35, x 40 respec-
tively. Figs 3, 4, posterior lateral and anterior lateral views, showing zigzag capillae noted in this
species only and often situated marginally. SEM x 20, x 15 respectively.

166 E. F. OWEN

The thickness varies according to the amount of secondary shell material deposited in older
individuals, but averages 6-9mm. It has a broad extensive triangular interarea and high
pseudodeltidium.

An average of 8-10 descending branches of the brachial apparatus are separated by thin
septae extending posteriorly from the shell margin.

A nearly perfect example of this species was figured by Backhaus (1959: pl. 4, figs 8, 9; pl. 5,
fig. 1) from the Middle Cenomanian, Essen Greensand, north Germany. Another Essen Green-
sand specimen is here figured (PI. 5, figs 18, 19) from the Davidson Collection, B5190.

Thecidiopsis essensis can be distinguished from the Hauterivian species T. tetragona and the
Maastrichtian species T. digitata by its constantly broad oval outline, acutely triangular and
extensive ventral interarea and short septum within the umbo of the pedicle valve. It has more
numerous and more parallel branches to its brachial apparatus than T. digitata, which it most
closely resembles, or than T. tetragona in which the branches are more lobate.

T. essensis has not been recorded from any horizon within the Cretaceous of Great Britain.
A specimen described, but not figured, by d’Archiac (1847: 350) from the Tourtia of Belgium as
Thecidea digitata Sowerby is probably T. essensis (Roemer).

Nekvasilova (1966: pl. 1, figs 1-6; 1974b: pl. 2, figs 3-6) described specimens from the Lower
Cretaceous of Stramberk, Czechoslovakia, as Thecidea cf. tetragona (Roemer); they have the
broad lobate brachial apparatus typical of T. tetragona, but bear some resemblance to T.
essensis in the width of valves and in the number and arrangement of the branches of the
brachial apparatus.

Faunal distribution

The distribution of brachiopod species of Boreal and Tethyan origin has been the subject of
much discussion since Ager (1971), Middlemiss (1975) and Owen (in Dieni, Middlemiss & Owen
1973) demonstrated the cosmopolitan nature of some of the forms previously considered to
have been confined to special ecological provinces. The Lower Chalk provides a good example,
amongst many, of faunal groups thought to have originated in these two major realms.

The adaptability of brachiopods to changes within the environment makes delimiting exact
ecosystems within such a community a very difficult task. It is fortunate, therefore, that the
universal transgressive Cenomanian sea provided relatively stable conditions, with few changes
in depth, so the resulting facies are limited. Ager (1965: 145) stressed the point that this stage
within the Mesozoic probably provides the best example of a stable, comparatively shallow-
water environment.

Within the broadly-defined Cenomanian are deposits which indicate areas of comparatively
deeper water or quieter conditions, which support their own peculiar brachiopod faunas. In
such places we see faunas somewhat analogous to Ager’s “Alpine Group’, but having closer
links with the overall fauna than the isolated ‘Alpine Group’ of species maintained with their
contemporaneous adjacent faunas.

In 1978 I pointed out, in a general review of brachiopod species within the Cenomanian of
northwestern and central Europe, that each ecological group within its own particular litho-
logical unit appears to contain a fauna with a characteristic or dominant association together
with some wider-ranging species which tend to fall into two groups; one characterized by
rhynchonellids and costate terebratellids and the other by terebratulids and smooth terebratel-
lids. The present study supports this view and I believe that these groups can be traced across
Europe into north Germany and southern France while retaining their specific constitution. In
so doing I have taken into consideration lithological and major geographical differences. For
instance, in southeastern France, at Drap, near Nice, a collection of brachiopods has been
made from highly glauconitic sandy limestone and from more argillaceous deposits nearby. The
species from the glauconitic beds consists of large biplicate Terebratulidae similar in general
form to Moutonithyris dutempleana (d’Orbigny) from similarly highly glauconitic deposits in
Britain and northern France, while the fauna in the more argillaceous deposits is of smooth

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 167

Terebratulidae and small Rhynchonellidae—an assemblage like the one which characterizes the
more marly beds within the Cenomanian of the Weald.

In Britain certain patterns can be recognized. In the south, particularly in the coarse sandy
facies of the more western outcrops at Wilmington and Warminster, there is a fauna showing
some characteristics of the two main lithologically controlled assemblages. I consider these to
be marginal to their normal faunal distributions. Genera such as Orbirhynchia, Grasirhynchia
and Terebratulina are found with acutely plicate Terebratulidae, such as Boubeithyris diplo-
plicata and coarsely costate Cyclothyridinae, and the Terebratellidae Dereta and Terebrirostra.
In addition, species such as the small ornate Arcuatothyris arcuata (Roemer) and Capillithyris
squamosa (Mantell) are commonly found in the chalk facies of the Lower Chalk Basement Beds
in Wiltshire and Dorset or the Craie de Rouen, where they occur with smooth, strongly
biconvex Terebratulidae such as Concinnithyris obesa (J. Sowerby). A. arcuata and C. squamosa
appear to be tolerant species, the former occurring also in the glauconitic limestones of north
Germany and on the Normandy coast; the latter, along with C. disparilis, in the Chalk Base-
ment Beds and Craie de Rouen and also in the Chalk Marl and marly facies of Kent and
Sussex. C. disparilis is found again with the other two species in sandy facies of Bed C at
Humble Point and at The Pinnacles, south Devon.

Grasirhynchia grasiana (d’Orbigny) appears to have adapted to coarse sandy glauconitic
limestones, as well as to more finely marly deposits of the Middle Cenomanian of the Weald,
Isle of Wight, Wiltshire, Dorset and Devon. It is also found widely in western and northern
France, rarely in the hard marly beds of Brochterbeck, Tecklenberg, north Germany, and in
Poland.

Summarizing, I consider that the Cenomanian transgressive sea extended from eastern
Europe across Poland, Czechoslovakia, north Germany, Belgium, France and Britain, and that
within this region there were areas of unusual environments. Further studies of these areas and
their brachiopod faunas may shed light on the differences between these environments, and on
the distributional parameters of the brachiopods.

It is possible to draw a broad comparison between the faunal composition of assemblages
from Essen, north Germany, the Arnager Greensand of Bornholm, Denmark, the Tourtia of
Belgium and the more argillaceous Cenomanian Sands of Wilmington, south Devon. Each of
these areas appears to represent an individual facies containing elements of both the marly bed
faunas and the harder, coarse sandy bed faunal elements discussed previously.

The more compact marly chalks and marls which occur within the same area seem to have
supported a fauna living on a softer substrate and perhaps in deeper water. This is reflected in
the composition of the assemblages which are characterized by smooth-shelled terebratulids
and finer costate rhynchonellids. The absence of any costate terebratellids is also characteristic
of quieter and deeper conditions.

Acknowledgements

I would like to express my thanks to colleagues at the British Museum (Natural History), in particular Dr
Howard Brunton, Dr Hugh Owen, Dr Robin Cocks, Mr Adrian Rissoné, Miss Lois Cody, Dr Noel
Morris, Dr Paul Taylor and Dr Richard Jefferies; also to the staff of the Photographic Studio, and Mr
Don Claugher and Miss Susan Barnes of the Electron Microscope Unit. Thanks are also due to Christo-
pher Wood, and Adrian Morter of the British Geological Survey, to Dr D. Price of the Sedgwick
Museum, Cambridge, to Dr Phil Drummond and Dr Rory Mortimore of Brighton Polytechnic, to Mr
Felix Whitham of Hull, to Dr Andrew Gale and Dr Keith Holdaway, to Mr Simon Hurt, Eastbourne, to
Dr Annie Dhondt, Belgium, Dr Eduard Lanterno, Museum d’Histoire Naturelle, Geneva, Dr Ewa Popiel-
Barczyk, Museum Ziemi, Warsaw, and Dr Olga Nekvasilova, Prague. My special thanks go to Dr Jake
Hancock, Imperial College, London, Dr F. A. Middlemiss, Queen Mary College, Dr W. J. Kennedy and
Dr C. W. Wright for much sound advice. Lastly, I wish to thank Dr John Ferguson and other members of
staff of Imperial College, for kindness and help.

168 E. F. OWEN

References

Ager, D. V. 1959. The classification of the Mesozoic Rhynchonelloidea. J. Paleont., Tulsa, 33: 324-332,
pl. 49.

—— 1965. The adaptation of Mesozoic brachiopods to different environments. Palaeogeogr. Palaeoclimat.
Palaeoecol., Amsterdam, 1: 143-172.

—— 1971. Space and time in brachiopod history. In Middlemiss, F. A., Rawson, P. F. & Newall, G. (eds),
Faunal provinces in Space and Time. Geol. J. Spec. Issue, Liverpool, 4: 95—110.

Allan, R. 1940. A revision of the Classification of the Terebratelloid Brachiopoda. Rec. Canterbury Mus.,
Christchurch, N.Z., 4 (6): 267-275. '

Amedro, F., Damotte, R., Manivit, H., Robaszynski, F. & Sornay, J. 1978. Echelles biostratigraphiques
dans le Cénomanien du Boulonnais (macro-micronanno fossiles). Geol. mediterr., Aix-en-Provence, 5 (1):
5-18.

, Dejonghe, L., Depuis, C. & Robaszynski, F. 1976. Les falaises crayeuses du Boulonnais: litho-
stratigraphie et repéres biostratigraphiques de l’Aptien au Senonien. C. r. somm. Seanc. Soc. geol. Fr.,
Paris, 1976 (3): 91—94.

Archiac, A. d’ 1847. Rapport sur les fossiles du Tourtia. Mem. Soc. géol. Fr., Paris, (2) 2: 291-351, pls
13-25.

Backhaus, E. 1959. Monographie der cretacischen Thecideidae (Brach.). Mitt. geol. StInst. Hamb., 28:
5—90, pls 1-7, 14 figs. :

Barrois, C. 1875. La zone a Belemnites plenus. Etude sur le Cenomanien et le Turonien du bassin de Paris.
Annls Soc. geol. N., Lille, 2: 146-193.

1876. Recherches sur le terrain crétace supérieur de |’Angleterre et de l’Irelande. Mem. Soc. geol. N..,
Lille, 1: 1-227, 3 pls, 15 figs.

Beecher, C. E. 1893. Revision of the families of loop-bearing Brachiopoda. Trans. Conn. Acad. Arts Sci.,
New Haven, 9 (2): 376-399, 3 pls.

Biernat, G. & Popiel-Barczyk, E. 1969. E. Panow—Materialy do znajomosci Ramienionogow z Gornej
Kredy okolic Krakowa [Contributions to the knowledge of the Brachiopoda from the Upper Creta-
ceous of the Krakow district]. Roczn. pol. Tow. geol., Krakow, 39 (4): 555-608, 3 pls, 2 figs.

Bigot, A. 1893. Sur quelques brachiopodes fossiles de Normandie. Bull. Lab. Geol. Fac. Sci. Caen, 2 (3):
113-120, 1 pl.

Bosworth, T. O. 1906. The zones of the Lower Chalk. Geol. Mag., London, 3 (5): 412-418.

Bower, C. R. & Farmery, J. R. 1910. The zones of the Lower Chalk of Lincolnshire. Proc. Geol. Ass.,
London, 21: 333-365.

Brocchi, G. 1814. Conchiologia fossile Subapennina con Osservazioni geologiche sugli Apennini e sul suolo
adiacente. 2 vols. 712 pp., atlas 15 pls. Milan.

Brogniart, A. 1822. In Cuvier, G. & Brongniart, A., Description geologique des environs de Paris. iv + 428
pp., 11 pls. Paris.

Bronn, H. G. 1835-37. Lethaea Geognostica, oder Abbildungen und Beschreibungen der fur die
Gebirgs-Formationen bezeichnendsten V ersteinerungen. 2 vols, atlas 47 pls. Stuttgart.

Bruguiere, J. G. 1797. Tableau Encyclopédique et Methodique des trois regnes de la Nature 2 (Vers,
Coquilles, Mollusques et Polypiéres): pls 190-286. Paris.

Buch, L. von 1838. Essai d’une classification et d’une description des Terébratules. Mem Soc. géol. Fr.,
Paris, (1) 3: 105—238, pls 13-20.

Carter, D. J. & Hart, M. B. 1977. Aspects of mid-Cretaceous stratigraphical micropalaeontology. Bull. Br.
Mus. nat. Hist., London, (Geol.) 29 (1): 1-135, 53 figs.

Casey, R. 1961. The stratigraphical palaeontology of the Lower Greensand. Palaeontology, London, 3:
487-621, pls 77-84.

Childs, A. 1959. Upper Jurassic rhynchonellid brachiopods from Northwestern Europe. Bull. Br. Mus. nat.
Hist., London, (Geol. Suppl.) 6: 1-119, pls 1-12, 40 figs.

Clerc, M. & Favre, J. 1910-18. Catalogue illustre de la Collection Lamarck. Premiére partie, Fossiles. 117
pls. Geneva.

Cooper, G. A. 1983. The Terebratulacea (Brachiopoda), Triassic to Recent: A study of the brachidia
(loops). Smithson. Contr. Paleobiol., Washington, 50: 1-291, pls 1-77, 17 figs, 86 tables.

Cox, M. M. 1967. See Pinder, M. M.

& Middlemiss, F. A. 1978. Terebratulacea from the Cretaceous Shenley Limestone. Palaeontology,
London, 21 (2): 411-441, pls 40-42.

Dagys, A. S. 1968. [Jurassic and early Cretaceous brachiopods of northern Siberia.] Trudy Inst. Geol.
Geofiz. sib. Otd., Novosibirsk, 41: 1-167, figs, tab.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 169

Dall, W. H. 180. A revision of the Terebratulidae and Lingulidae. Am. J. Conch., Philadelphia, 6: 88-168,
pls 6-8.

1900. Some names which must be discarded. Nautilus, Philad., 14 (4): 44-45.

Dames, W. 1874. Ueber Diluvial-Geschiebe cenomanen Alters. Z. dt. geol. Ges., Berlin, 27: 61-773, pl. 21
fig. 5.

Davidson, T. 1850. Notes on an examination of Lamarck’s species of fossil Terebratulidae. Ann. Mag. nat.
Hist., London, (2) 5: 434450, 3 pls.

1852-55. A Monograph of British Cretaceous Brachiopoda. 117 pp., 12 pls. Palaeontogr. Soc.

(Monogr.), London.

1874. A Monograph of the fossil Brachiopoda, 4 (1); supplement to the Recent, Tertiary and

Cretaceous Species. 72 pp., 8 pls. Palaeontogr. Soc. (Monogr.), London.

& Morris, J. 1847. Description of some species of Brachiopoda. Ann. Mag. nat. Hist., London, (1) 20:
250-258, 2 pls.

Defrance, M. J. L. 1828. Terebratula deformis Lamarck. In Levrault, F. G. (ed.), Dictionnaire des Sciences
Naturelles 53: 160. Strasbourg & Paris.

Deicke, H. 1878. Beitrage zur Kenntniss der geognostischen und paldontologischen Beschaffenheit der
unteren Ruhrgegend, 2. Die Brachiopoden der Tourtia von Miilheim a.d. Ruhr. Jber. Realschule 1.
Miilheim a.d. Ruhr 25. 28 pp., 1 pl.

Dibley, G. E. 1918. Additional notes on the Chalk of the Medway Valley, Gravesend, West Kent,
North-east Surrey, and Grays (Essex). Proc. Geol. Ass., London, 29: 68-105, pls 7-9, figs 8-10.

Dieni, I., Middlemiss, F. A. & Owen, E. F. 1975. The Lower Cretaceous Brachiopods of east-central
Sardinia. Boll. Soc. paleont. ital., Modena, 12 (2): 166-216, pls 32-38, figs 1-18.

Donovan, D. T. 1949. Observations on the Mesozoic rocks of Geographical Society @, East Greenland.
Medadr Gronland, Copenhagen, 149 (5): 1-13.

Drummond, P. V. O. 1970. The Mid-Dorset Swell. Evidence of Albian—Cenomanian movements in
Wessex. Proc. Geol. Ass., London, 81 (4): 679-716.

Elliott, G. F. 1938. A London Clay Brachiopod. Proc. Geol. Ass., London, 49: 128-134, figs.

1959. Six new genera of Mesozoic Brachiopoda. Geol. Mag., Hertford, 96: 146-148.

1965. Mesozoic Terebratellacea: discussion. in Muir-Wood, H. M. et al. (q.v.). In Moore, R. C. (ed.),
Treatise on Invertebrate Paleontology H (Brachiopoda): H818—H819. Lawrence, Kansas.

Ernst, G. & Schmid, F. 1979. Multistratigraphische Untersuchungen in der Oberkreide des Raumes
Braunschweig-Hannover. In Wiedmann, J. (ed.), Aspekte der Kreide Europas: 11-46. Stuttgart, Beitrage
zum 1. Symposium Deutsche Kreide (Int. Un. geol. Sci. (A) 6).

Etheridge, R. 1881. Palaeontology. In Penning, W. H. & Jukes-Browne, A. J., The Geology of the
neighbourhood of Cambridge: 147, pl. 3. Mem. geol. Surv. U.K., London.

Favre, J. 1918. See Clerc, M. & Favre, J.

Fletcher, T. P. 1977. Lithostratigraphy of the Chalk (Ulster White Limestone Formation) in Northern
Ireland. Rep. Inst. geol. Sci., London, 77/24. 33 pp.

& Wood, C. J. 1983. Geology of the country around Carrickfergus and Bangor. Mem. geol. Surv.
Nth. Ire., Belfast, sheet 29: 44-54, figs 14-18.

Frebold, H. 1935. Marine Aptien von der Koldeway Insel (Nordliches Ostgr6nland). Meddr Gronland,
Copenhagen, 95 (4): 1-112, pls 1-8.

Gallois, R. W. & Morter, A. 1978. Appendix 1. The Upper Jurassic and Cretaceous stratigraphy of IGS
boreholes 72/77 and 72/78. Rep. Inst. geol. Sci., London, 78/18: 24-31.

Gaspard, D. 1971. Variabiliteé morphologique d'une population de Sellithyris (Térébratules de la région du
Mans—Cénomanien supérieur). Notes Contr. Cent. Etud. Rech. Paleont. biostratigr., Orsay, 1: 1-12,
4 pls.

1976. Caractéres Morphologiques et Microstructure de ‘Terebratula’ dutempleana d’Orbigny. Annls
Paleont., Paris, (Invert.) 62: 123-138, 5 pls.

Gaster, C. T. A. 1929. Chalk zones in the neighbourhood of Shoreham, Brighton and Newhaven, Sussex.
Proc. Geol. Ass., London, 40: 328-340, pl. 27.

1937-51. The stratigraphy of the Chalk of Sussex. 1: West Central area, Arun Gap to valley of the
Adur. Proc. Geol. Ass., London, 48: 356-373 (1937). 2: Eastern area, Seaford, Cuckmere valley and
Eastbourne. Loc. cit. 50: 510-526 (1939). 3: Western area, Arun Gap to the Hampshire boundary. Loc.
cit. 55: 173-188 (1944). 4: East central area, between the valley of the Adur and Seaford. Loc. cit. 62:
31-64 (1951). With zonal maps.

Geinitz, H. B. 1872. Das Elthalgebirge in Sachsen. Palaeontographica, Stuttgart, 20: 147—207, pls 36-45.

Gray, J. E. 1840. Synopsis of the Contents of the British Museum. 42nd edn. 370 pp. London.

1848. On the Arrangement of the Brachiopoda. Ann. Mag. nat. Hist., London, (2) 2: 435-440.

5)

170 E. F. OWEN

Hagenow, F. von 1842. Monographie der Rugen’schen Kreide-Versteinerungen III. Abt. Mollusken. Neues
Jb. Miner. Geogn. Geol. Petrefakt., Stuttgart, 1842: 528-575, pl. 9.

Hamblin, R. J. O. & Wood, C. J. 1976. The Cretaceous (Albian—Cenomanian) stratigraphy of the Haldon
Hills, south Devon, England. Newsl. Stratigr., Berlin & Stuttgart, 4 (3): 135-149, 2 figs, 1 tab.

Hancock, J. M. 1961. The Cretaceous system in Northern Ireland. Q. JI geol. Soc. Lond., 117: 11-36.

Hofman, E. 1869. Materials for the Geology of Russia 1.99 pp., 19 pls. St Petersberg.

Huxley, T. 1869. An introduction to the classification of animals. 147 pp., 47 figs. London.

Jarvis, I. & Woodroof, P. B. 1984. Stratigraphy of the Cenomanian and basal Turonian (Upper Creta-
ceous) between Branscombe and Seaton, SE Devon, England. Proc. Geol. Ass., London, 93 (3): 193-215,
figs 1-6, 1 tab.

Jeans, C. V. 1968. The origin of the Montmorillonite of the European Chalk with special reference to the
Lower Chalk of England. Clay Miner., Oxford, 7: 311-329, figs & tables.

1973. The Market Weighton structure, tectonics, sedimentation and diagenesis during the Creta-
ceous. Proc. Yorks. geol. Soc., Leeds, 39: 409-444.

Jefferies, R. P. S. 1962. The palaeoecology of the Actinocamax plenus Subzone (Lowest Turonian) of the
Anglo-Paris Basin. Palaeontology, London, 4: 609-647.

1963. The stratigraphy of the Actinocamax plenus Subzone (Turonian) in the Anglo-Paris Basin.
Proc. Geol. Ass., London, 74: 1—34.

Juignet, P. (1974). La transgression Cretace sur la bordure orientale du Massif Armoricaine. Aptien, Albien,
Cenomanien de Normandie et du Maine. Le stratotype du Cénomanien. 810 pp., 28 pls. Thése, Univ.
Caen.

& Kennedy, W. J. 1976. Faunes d’ammonites et biostratigraphie comparée du Cénomanian du

nord-ouest de la France (Normandie) et du sud de l’Angleterre. Bull. Soc. geol. Normandie, Le Havre,

63: 1-193, pls 1-34.

, & Wright, C. W. 1973. La limite Cénomanien-Turonien dans la région du Mans (Sarthe):
stratigraphie et paléontologie. Annls Paléont., Paris, (Invert.) 59: 209-242.

Jukes-Browne, A. J. 1900. Cretaceous Rocks of Britain, 1. Gault and Upper Greensand of England.
x + 499 pp., 5 pls. Mem. geol. Surv. U.K., London. See Jukes-Browne & Hill 1903-04.

& Hill, W. 1896. A delimitation of the Cenomanian. Q. JI geol. Soc. Lond., 52: 99-177, pl. 5, 3 tables.

1903-04. Cretaceous Rocks of Britain, 2. Lower and Middle Chalk of England. xiii + 568 pp.,
8pls. Mem. geol. Surv. U.K., London. See Jukes-Browne 1900.

Katz, J. I. 1974. In Krymgol'’ts, G. Y. (ed.), [Atlas of the Upper Cretaceous fauna of the Don Basin]. 640 pp.,
128 pls. Moscow [In Russian].

Kennedy, W. J. 1969. Correlation of the Lower Chalk of south-east England. Proc. Geol. Ass., London, 80:
459-560, pls 15-21, 15 figs.

1970. Correlation of the uppermost Albian and Cenomanian of south-west England. Proc. Geol. Ass.,

London, 81: 613-677, 17 figs.

& Hancock, J. M. 1970. Ammonites of the genus Acanthoceras from the Cenomanian of Rouen,
France. Palaeontology, London, 13: 462-490, pls 88-97.

King, W. 1850. A monograph of the Permian fossils of England, 3. xxxvii + 258 pp., 29 pls. Palaeontogr.
Soc. (Monogr.), London.

Lamarck, J. B. P. 1819. Histoire naturelles des animaux sans vertebres (1st. edn), 6 (6). 343 pp. Paris.

Lamplugh, G. W. & Walker, J. F. 1903. On a fossiliferous band at the top of the Lower Greensand near
Leighton Buzzard (Bedfordshire). Q. JI geol. Soc. Lond., 59: 234-265, 3 pls.

Lang, W. D. 1904. The zone of Hoplites interruptus (Bruguiére) at Black Ven, Charmouth. Geol. Mag.,
London, (5) 1: 124-231, 4 figs.

Lehner, L. 1937. Fauna und Flora der frankischen albiiberdeckenden Kreide. 2: Fauna. Palaeonto-
graphica, Stuttgart, A87: 158—230, pls 16-19, 3 figs.

M‘Coy, F. 1844. A synopsis of the characters of the Carboniferous Limestone fossils of Ireland. viii + 207
pp., 29 pls. Dublin.

McGugan, A. 1957. Upper Cretaceous foraminifera from Northern Ireland. J. Paleont., Tulsa, 31: 329-
348, pls 30-35.

Makridin, V. P. 1955. [Some Jurassic rhynchonellids of the European part of the USSR.] Zap. geol. Fak.
Kharkov. Gos. Univ., 12: 81—91 [In Russian].

Mantell, G. A. 1822. Fossils of the South Downs or illustrations of the Geology of Sussex. xvii + 327 pp.,
42 pls. London.

Menke, C. T. 1828. Synopsis methodica molluscorum generum omnium et specierum earum quae in Museo
Menkeano adservantur. 91 pp. Pyrmonti.

Meyer, C. J. A. 1874. On the Cretaceous Rocks of Beer Head and the adjacent Cliff-sections, and on the

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 171

Relative Horizons therein of the Warminster and Blackdown Fossiliferous Deposits. Q. JI geol. Soc.
Lond., 30: 369-393. [See also Geol. Mag., London, (2) 1: 375-377].

Middlemiss, F. A. 1959. English Aptian Terebratulidae. Palaeontology, London, 2 (1): 94-142, pls 15-18.

1975. In Dieni, I., Middlemiss, F. A. & Owen, E. F., q.v.

1976. Lower Cretaceous Terebratulidina of Northern England and Germany and their geological

background. Geol. Jb., Hannover, (A) 30: 21—104, pls 1-11.

1978. The genus Platythyris (Brachiopoda) and its relationship to the Pygopidae. Palaont. Z.,
Stuttgart, 52 (1/2): 28-46.

Morter, A. A. & Wood, C. J. 1983. The biostratigraphy of Upper Albian-Lower Cenomanian Aucellina in
Europe. Zitteliana, Munich, 10: 515-529.

Muir-Wood, H. M., Stehli, F. G., Elliott, G. F. & Hatai, K. 1965. Terebratulida. In Moore, R. C. (ed.),
Treatise on Invertebrate Paleontology H (Brachiopoda): H728—H857. Lawrence, Kansas.

Nekvasilova, O. 1966. A Thecidean brachiopod new to the Lower Cretaceous of Czechoslovakia. Vést.
ustred. Ust. geol., Prague, 41 (3): 193-194, 2 pls.

1974. The genus Thecidioposis (Brachiopoda) from the Lower Cretaceous of Stramberk (Czechoslo-

vakia). Cas. Miner. Geol., Prague, 19 (3): 239-244, pls. 1, 2, 1 fig.

1983. The genus Argyrotheca (Brachiopoda) from the Bohemian Cretaceous Basin (Czechoslovakia).
Cas. Miner. Geol., Prague, 28 (1): 23-30, pls 1-4, 2 figs.

Oehlert, D. P. 1887. Appendice. Brachiopodes. In Fischer, P., Manuel de Conchyliologie et de Paleontol-
ogie Conchyliologique, ou Histoire Naturelle des Mollusques vivants et fossiles: 1189-1334. Paris.

Orbigny, A. d’. 1847—S1. Terrains crétacés. Paleontologie frangaise 4: 1-390, pls. 490-599. Paris.

1852. Cours Elementaire de Paleontologie et de Geologie Stratigraphiques 2 (2): 383—847. Paris.

Owen, E. F. 1962. The brachiopod genus Cyclothyris. Bull. Br. Mus. nat. Hist., London, (Geol.) 7 (2):
39-63, 2 pls, 10 figs.

— 1963. The brachiopod genus Modestella in the Lower Cretaceous of Great Britain. Ann. Mag. nat.
Hist., London, (13) 6: 199-203, 1 pl., 3 figs.

— 1968. A further study of some Cretaceous rhynchonelloid brachiopods. Bull. Indian Geolists Ass.,
Chandigarh, 1: 17-32, 1 pl., 3 figs.

— 1970. A revision of the brachiopod subfamily Kingeninae Elliott. Bull. Br. Mus. nat. Hist., London,
(Geol.) 19 (2): 27-83, pls 1-14, 17 figs.

—— 1975. In Dieni, I., Middlemiss, F. A. & Owen, E. F., q.v.

— 1977. Evolutionary trends in some Mesozoic Terebratellacea. Bull. Br. Mus. nat. Hist., London,
(Geol.) 28 (3): 205-253, 3 pls, 29 figs.

1978. The distribution of Brachiopods within the Cenomanian of northwest and central Europe.
Geol. méditerr., Aix-en-Provence, 5 (1): 147-154, 4 figs.

Panow, E. 1969. (posthumous publication: see Biernat & Popeil-Barczyk). ;

Parent, H. 1893. Sur l’existence du Gault entre les Ardennes et le Bas Boulonnais (Etude du Gault du
Cenomanien de Artois). Annls Soc. geol. N., Lille, 21: 205—247.

Pauca, M. & Patrulius, D. 1960. Contribution a l’étude paléontologique des depdts Albiens de Giurgiu
(Vallée du Danube). Studii Cerc. Geol., Bucharest, 5 (1): 85-100, 6 pls, figs.

Pettitt, N. 1949-54. A monograph of the Rhynchonellidae of the British Chalk. I: i-iv, 1-26, pls 1-2
(1949). II: 27-52, pl. 3 (1954). Palaeontogr. Soc. (Monogr.), London.

Pinder, M. M. (1967). Studies of some English Albian Terebratulidae. Text & Plates. Ph.D. Thesis, Univ.
London (unpublished).

Popiel-Barczyk, E. 1972. Albian—Cenomanian brachiopods from the environs of Annopol on the Vistula
with some remarks on related species from Cracow, Poland. Pr. Muz. Ziemi, Warsaw, 20: 119-149, pls
1-4, 12 figs.

1977. A further study of Albian—Cenomanian brachiopods from the environs of Annopol on the

Vistula with some remarks on related species from the Cracow region, Poland. Pr. Muz. Ziemi,

Warsaw, 26: 25-54, pls 1-7, 20 figs.

& Smirnova, T. N. 1978. [New genus of Megathyrid (Brachiopoda) from the Upper Cretaceous of
Poland.] Paleont. Zh., Moscow, 1978 (3): 134-137.

Price, F. G. H. 1877. On the beds between the Gault and Upper Chalk near Folkestone. Q. JI geol. Soc.
Lond., 33: 431-448.

Quenstedt, F. A. 1868-71. Die Brachiopoden. In: Petrefactenkunde Deutschlands, (1) 2. iv + 748 pp., atlas
pls 37-61. Leipzig.

1882. Handbuch der Petrefaktenkunde. viii + 1239 pp. Tubingen.

Rayn, J. P. J. 1916. Kridttaflejringerne paa Borholms Sydvestkyst og deres Fauna. I. Cenomenat. Danm.
geol. Unders., Copenhagen, (2) 30: 1-39, pls 1—5.

172 E. F. OWEN

— 1925. Det cenomane Basalkonglomerat paa Bornholm. [Sur le conglomerat de base du Cénomanien
de Vile de Bornholm.] Danm. geol. Unders., Copenhagen, (2) 42: 1-56, 4 pls.

Rawson, P. F. et al. 1978. A correlation of Cretaceous rocks in the British Isles. Spec. Rep. geol. Soc. Lond.
9: 1-70.

Reid, R. E. H. 1958a. The Upper Cretaceous Hexactinellidae. Part I. 46 pp., 24 figs. Palaeontogr. Soc.
(Monogr.), London.

— 1958b. Remarks on the Upper Cretaceous Hexactinellidae of County Antrim. Ir. Nat. J., Belfast, 12:
236-243, 261-269.

— 1971. The Cretaceous rocks of north-east Ireland. Ir. Nat. J., Belfast, 17: 105—129.

Roemer, F. A. 1840-41. Die Versteinerungen des norddeutschen Kreidegebirges (1): 1-48, pls 1-7 (1840); (2)
49-145, pls 8-16 (1841). Hannover.

—— 1854. Die Kreidebildungen Westphalens. Ein geognostische Monographie. Verh. naturh. Ver. preuss.
Rheinl., Bonn, 11 (4): 29-180, map.

Rollier, L. 1911. Fossiles nouveaux ou peu connus des terrains secondaires du Jura et des contrées
environants, 1. Abh. Schweiz. palaont. Ges., Geneva, 37 (1): 1-31, pls 1-5.

Rzhonsnitzkaya, M. A. 1959. [Systematics of the Rhynchonellids]. Paleont. Zh., Moscow, 1959 (1): 25-36.

Sahni, M. R. 1929. A monograph of the Terebratulidae of the British Chalk. 62 pp., 10 pls. Palaeontogr.
Soc. (Monogr.), London.

1958. Supplement to a monograph of the Terebratulidae of the British Chalk. Monogr. palaeont. Soc.
India, Lucknow, 1: 1—22, 6 pls.

Schloenbach, G. J. C. U. 1866a. Mittheilungen an Professor H. B. Geinitz. Neues Jb. Miner. Geol. Palaont.,
Stuttgart, 1866: 574-575.

—— 1866b. Beitrage zur Palaeontologie der Jura- und Kreide-Formation im nordwestlichen Deutsch-
land. II. Kritische Studien ber Kreide-Brachiopoden. Palaeontographica, Cassel, 13: 267-332, 3 pls.

1867. Uber die Brachiopoden der Norddeutschen Cenoman-Bildungen. Geogn.-palaont. Beitr.,
Munich, 1 (3): 405—S06, pls 21—23.

Schlotheim, E. F. von 1813. Beitrage zur Naturgeschichte der Versteinerungen in geognostischer Hinsicht.
Taschenb. ges. Miner., Frankfurt a.M., 7: 1-134.

1820-23. Die Petrefactenkunde euiinen jetzigen Standpunkte durch die Beschreibung seiner Sammlung
versteinerter und fossilier Uberreste des Thier- und Pflanzenreichs der Vorwelt erlautert. \xii + 437 pp., 54
pls. Gotha.

Schuchert, C. 1900. Brachiopoda. In Zittel, K. A. von (Eastman, C. R., transl. & ed.), Textbook of
Palaeontology 1 (1st edn): 291—343, figs. London.

Selwood, E. B. et al. 1984. Geology of the country around Newton Abbot. 212 pp. Mem. Br. geol. Surv.,
London, sheet 339 (n.s.).

Smirnova, T. N. 1972. [Early Cretaceous brachiopods of the Crimea and the northern Caucasus]. 143 pp., 13
pls. Moscow, Izdatel’stvo ‘Nauka’ [In Russian]. (Publ. version of thesis, Moscow State University,
1963).

Smith, A. B., Paul, C. R. C., Gale, A. S. & Donovan, S. K. 1988. Cenomanian and Lower Turonian
echinoderms from Wilmington, south-east Devon, England. Bull. Br. Mus. nat. Hist., London, (Geol.)
42: 1-245.

Smith, W. E. 1957a. The Cenomanian Limestone of the Beer district, South Devon. Proc. Geol. Ass.,
London, 68: 115-133, pls 4-5, 5 figs.

— 1957b. Summer Field Meeting in South Devon. Proc. Geol. Ass., London, 68: 136-152.

— 1961. The Cenomanian deposits of south-east Devonshire. Proc. Geol. Ass., London, 72: 91-133, pls
4-6, 8 figs.

Sowerby, J. 1812-22. The Mineral Conchology of Great Britain 1: 1-234, pls 1-102 (1812-15); 2: 1-251, pls
103-203 (1815-18); 3: 1-184, pls 204-306 (1818-21); 4 (part): 1-106, pls 307—377 (1821-22). London.

Sowerby, J. de C. 1822-46. The Mineral Conchology of Great Britain 4 (part): 107-160, pls 378-406
(1822-23); 5: 1-116, pls 407-503 (1823-25); 6: 1-236, pls 504-609 (1826-29); 7: 1-80, pls 610-648
(1840-46). London.

1836. In Fitton, W. H., Observations on some of the strata between the Chalk and the Oxford Oolite
in the south-east of England. Trans. geol. Soc. Lond., (2) 4: 103-389, pls 11—23.

Tate, R. 1880. On the Australian Tertiary Palliobranchs. Trans. Proc. r. Soc. South Austr., Adelaide, 3:
140-170, pls 7-11.

Thomel, G. 1965. Zonéostratigraphie et Paléobiogéographie du Cénomanien du sud-est de la France. C,r.
Congr. Socs sav. Paris Sect. Sci. 90 (Nice) (2): 127-154.

Tiessen, E. 1895. Die subhercyne Tourtia und ihre Brachiopoden- und Mollusken-fauna. Z. dt. geol. Ges.,
Berlin, 47: 423-533, pls 7-8.

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 173

Topley, W. 1875. The Geology of the Weald (parts of the counties of Kent, Surrey, Sussex and Hants).
xiv + 503 pp., 2 maps. Mem. geol. Surv. U.K., London.

Valenciennes see Lamarck.

Wahlenberg, G. 1821 (1819). Petrificata telluris Svecanae examinata. Nova Acta R. Soc. Scient. upsal. 8:
1-116, pls 14.

Walker, J. F. 1903. In Lamplugh, G. W. & Walker, J. F., q.v.

Wanderer, K. 1909. Die wichtigsten Tierversteinerungen aus der Kreide des Konigreiches Sachsen.
xxxxii + 80 pp., 12 pls. Jena.

Weidmann, J. 1979. Die Ammoniten der NW-deutschen, Regensberger und Ostalpinen Oberkreide im
Vergleich mit den Oberkreidefaunen des westlichen Mediterrangebeites. In Weidmann, J. (ed.), Aspekte
der Kreide Europas: 335-350. Stuttgart, Beitrage zum 1. Symposium Deutsche Kreide (Int. Un. geol.
Sci. (A) 6).

Wilson, V. et al. 1958. Geology of the country around Bridport and Yeovil. Mem. geol. Surv. U.K..,
London, 32: 230-235.

Wisniewska, M. 1932. Les Rhynchonellides du Jurassique supérieur de Pologne. Palaeont. pol., Warsaw, 2:
1-71, pls 1-6.

Wright, C. W. & Kennedy, W. J. 1981. The Ammonoidea of the Plenus Marls and the Middle Chalk.
148 pp., 32 pls. Palaeontogr. Soc. (Monogr.), London.

1984. The Ammonoidea of the Lower Chalk. Part 1. pp. 1-126, pls 1-40. Palaeontogr. Soc.

(Monogr.), London.
—— & Wright, E. V. 1942. The Chalk of the Yorkshire Wolds. Proc. Geol. Ass., London, 53: 112-127,

fig. 8.
Zareczny, S. 1874. O sredniém ogniwie warstw cenomanskich w Galicyi wschodniéj. Spraw. Kom. fizyogr.

Krakow, 8: 99-183, 2 pls.

Index

“Alpine Group’ 166 Cap Blanc Nez 79, 91, 95, 150
Annopol, Poland 91, 119 Cap de la Héve 80, 85, 109
Arnager Greensand, Bornholm 69, 167 Cassis, France 134, 138, 142

Cenomanian zones 71
Balcombe pit 69 Chalk Marl 142
Ballard Point 73, 142, 162 Chalk Rock 132
Barrington, Cambridge 96 Channel tunnel boreholes 70
Beachy Head, E. Sussex 69, 93 Chard, Somerset 85, 109, 130
Beddingham, E. Sussex 93 Chardstock, Devon 98-9, 109, 130, 142, 144, 165
Beer Head, Devon 98 Charente, France 80
Belfast Marls 79 Charing, Kent 92
Belgium 80 Cherry Hinton, Cambridge 148
Betchworth, Surrey 99 Chinnor, Oxford 150
Bornholm, Denmark 167 Chute Farm 91-2, 108, 114
Boubeithyris diploplicata sp. nov. 112-14 Clapham’s Limeworks, Louth 148
Boulonnais, France 79 Colinwell Sands 79
Bousse, France 80, 96, 103 Compton Bay 84, 91, 104, 109, 111
Bovey Lane, Devon 99, 102-3, 147 Céte Ste Catharine 96, 112, 136, 147
Briollay, France 89, 106 Cran d’Escalles 91, 95, 150
Brockterbeck, Germany 91, 144 Craie de Rouen 80, 128, 131, 134, 136, 138, 147,
Brunni Alp, Bernese Oberland 134 167
Bryozoansandstein 81 Culver Cliff, Isle of Wight 91, 116
Burham, Kent 51, 70 Cyclothyris formosa sp. nov. 85-6
Burrirhynchia devoniana sp. nov. 89-90 juigneti sp. nov. 86
Burwell, Cambridge 48, 95, 150 punfieldensis sp. nov. 85

Burwell, Lincolnshire 96
; ; Dereta pectita ballardensis subsp. nov. 162
Caistor, Lincolnshire 138 Deverill Hill 114, 150
Cambridge Greensand 122, 124, 127-8, 138, 147- Dover, Kent 103, 148, 150
8, 156 Drap, near Nice 142, 166

174 E. F. OWEN

Dresden, Germany 81
Dunscombe, Devon 162
Durdle Cove, Dorset 130

Eastbourne, E. Sussex 69, 103, 112, 150

Eastern outcrops 76

Eggardon Hill 128, 140, 147

Elsham, Lincolnshire 138

Essen, Germany 102, 144; Greensand 69, 81, 95,
101, 104, 166

Faunal Distribution 166

Feécamp, France 112

Fetcham Mill borehole 70

Floteberg, Germany 150

Folkestone, Kent 93, 95, 103, 144, 148, 154, 160,
162

Fulbourn, Cambridge 134, 138, 140, 142

‘Gault’ of Black Ven, Dorset 84
Glauconitic Marl 84

Glynde, E. Sussex 69, 140, 150
Goneville, France 104

Gore Cliff, Isle of Wight 91, 116
Grandpré, France 84
Greenhithe, Kent 138

‘Grizzle’ 107, 114

Grove Mill Pit 134, 142
‘Griinsand von Quedlinburg’ 154, 156
Gussignies, Belgium 80

Haldon Hills 74, 90

Hallington 92

Hamsey, E. Sussex 91-3, 100-1, 109, 142, 148,
150, 152, 154

Heacham, Norfolk 138

Head Ledge, Eastbourne 69

Heytesbury, Wiltshire 91—2

High Melcombe, Dorset 85

Hillington, Norfolk 138

Hilsconglomerat, Essen 69, 152

Hitchin, Hertfordshire 96

Hooken Cliff, Devon 98

Horn Hill, Beaminster 147

Horningsham, Wiltshire 142, 160

Hubbard’s Vale, Louth 150

Humberside 77

Humble Point, Devon 147, 167

Hunstanton, Norfolk 96, 101, 120, 122-4, 127-8,
144, 148

Island Magee Siltstone 79
Isle of Wight 84, 130, 136, 150, 154
Isleham 96

Julianka, Poland 116, 119

Koldeway Island, Greenland 124
Krakow, Poland 154

La Perriére, France 86
Langelsheim, Germany 81

Le Havre, France 90, 115, 150

Le Mans, France 80, 89, 106, 142
Leighton Buzzard 85, 90, 112
Lewis, E. Sussex 130

Liques, Pas de Calais 154

Little Beach, Devon 96, 99

London Clay, Sheppey 152

Louth, Lincolnshire 92, 116, 119-20, 140
Lower Pink Band 78

Lower Tertiary 147

Lydden Spout, Folkestone 148, 150

Maiden Newton 109

Marham 96

Market Weighton, Humberside 148, 152
Marnes a Ostracées 89

Mayet, France 89, 106

Melcombe Bingham 90, 162

Melton, S. Yorkshire 92; Carstone 112
Mere, Wiltshire 90

Merstham, Surrey 99

Méziéres sous Ballon 166

Montigny sur Roc 95

Morasheimer 81

Moutonithyris anglia sp. nov. 126-7

“Neanic Holotype’ 132
Newington’s Pit 69

Newtimber Place 150

Norfolk 76, 154

North Downs-inland exposures 72
North-east England 77

Northern France 79

Northern Ireland 78

Norton Bavant 128, 136

Norton Ferris 96

Octerville, France 80, 91

‘Orbirhynchia’ Beds 9

Orbirhynchia boussensis sp. nov. 103
wilmingtonensis sp. nov. 102-3; variation 101

Ovatathyris potternensis gen. et sp. nov. 106-12

Oxted, Surrey 138

‘Paradoxica’ Bed 77

Perte du Rhone 109, 111

Pinnacles, Devon 147, 167

Plenus Marls 78

Port des Barques, France 106, 109, 142
Potterne, Wiltshire 101, 107, 109, 111, 158
Punfield, Swanage 73, 85, 150, 160, 162

Rectithyris wrightorum sp. nov. 114-16

Red Rock, Hunstanton 76-7, 126

Reigate, Surrey 92

Regensburg, Germany 81

Rocken End, Isle of Wight 91, 104, 112, 116, 120,
150

CENOMANIAN BRACHIOPODS FROM LOWER CHALK OF BRITAIN AND NORTH EUROPE 7/5)

Rodmill Cement Works 69 Terebratulina etheridgei nom. nov. 147
Rouen, France 80, 165 imbricata sp. nov. 148, 150
Rougefort, Pas de Calais 91, 154 protostriatula sp. nov. 150, 152

Rifle Butts Pit 148, 152 Toller Fratrum, Porcorum 147

Rye Hill Farm 114 Totternhoe Stone 76-8, 95-6

Tournai, Belgium 80, 95, 101, 127, 142, 144
Tourtia 80-1, 144, 148, 150

Tropeothyris vectis sp. nov. 116-20
Sable de Bousse 102

Sables du Perche 80, 89, 104 Ulster White Limestone 79

Sables et Grés du Mans 42, 89, 104 Undercliff, Isle of Wight 120

St Aybert, France 124

St Jouin, France 86, 109 Vimoutiers, France 91, 109, 112

St Lawrence, Isle of Wight 150

Shapwick, Dorset 128 Warminster, Wiltshire 85, 90-1, 107-8, 111, 142,
Snowdon Hill, Chard 84, 138, 147 150, 156, 158, 167; Greensand 131

South Ferriby, Humberside 128 White Nothe, Dorset 130

Southerham Grey Pit 69, 93, 133 Whitecliff, Isle of Wight 90

South-western outcrops 73 Wilmington, Devon 75, 80, 89-90, 101-2, 115,
Speeton Cliff, N. Yorkshire 138, 140 142, 147, 150, 167

Storridge Hill 128 Woody Bay, Isle of Wight 91

Swaffham Prior, Cambridge 96 Wunstorf, Germany 81, 91

Accepted for publication 9 November 1986

- 7 ee es tye ty CA ao ya

Bulletin of the British Museum (Natural History)
Geology Series

Most earlier Geology Bulletins are still in print. A full list of available titles can be obtained from Publications Sales
(address inside front cover).

Vol. 31 No. 1 Foraminifera of the Togopi Formation, eastern Sabah, Malaysia. J. E. Whittaker & R. L. Hodgkinson.
1979. Pp. 1-120, 10 plates, 71 figs. £14.00
Vol. 31 No. 2 Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Gaudryceratidae.
W. J. Kennedy & H. C. Klinger. 1979. Pp. 121-173. £6.25
Vol. 31 No. 3 Benthic community organization in the Ludlow Series of the Welsh Borderland. R. Watkins. 1979. Pp.
175-279. £12.25
Vol. 31 No.4 The ammonites of the English Chalk Rock (Upper Turonian). C. W. Wright. 1979. Pp. 281-330. £6.50

Vol. 32 No.1 Miscellanea: Observations on Cycloclypeus—Provenance of Sivapithecus—Iranian Silurian
brachiopods—New English condylarths—Miocene sharks’ teeth—East African isopod—The Singa skull—

Carboniferous insects. 1979. Pp. 1-90. £10.50
Vol. 32 No. 2 Palaeoenvironments and correlations of the Carboniferous rocks in west Fermanagh, Ireland. C. H. C.
Brunton & T. R. Mason. 1979. Pp. 91-108, 6 figs, folded map. £4.00
Vol. 32 No.3 The Ordovician trilobite faunas of the Builth—Llandrindod Inlier, central Wales. Part III. C. P. Hughes.
1979. Pp. 109-181, 177 figs. £10.00
Vol. 32 No.4 The stratigraphy and brachiopods of the upper part of the type Caradoc of south Salop. J. M. Hurst.
1979. Pp. 183-304, 557 figs. £18.50

Vol. 33 No.1 An account of the Ordovician rocks of the Shelve Inlier in west Salop and part of north Powys. W. F.
Whittard, F. R. S. (Compiled by W. T. Dean). 1979. Pp. 1-69, 38 figs, frontispiece, coloured map, folded, in pocket.
(Map available separately for £1.00) £10.00

Vol. 33 No.2 Miscellanea: Lower Carboniferous microproblematicum—Miniature trilobite—Pleistocene bird
remains—English Eocene Hyracotherium—Salenia trisuranalis—Antarctic brachiopods—Diphyphyllum and Murchi-

son’s Russian corals—Lebanese amber Neuroptera. 1980. Pp. 71-164. £12.00
Vol. 33 No.3 The Caradoc faunal associations of the area between Bala and Dinas Mawddwy, north Wales. M. G.
Lockley. 1980. Pp. 165-235, 105 figs. £9.00
Vol. 33 No. 4 Fossil insects from the Bembridge Marls, Palaeogene of the Isle of Wight, southern England. E. A.
Jarzembowski. 1980. Pp. 237-293, 77 figs. £7.50
Vol. 33 No.5 The Yorkshire Jurassic fern Phlebopteris braunii (Goeppert) and its reference to Matonia R.Br. T. M.
Harris. 1980. Pp. 295-311, 20 figs. £2.75

Vol. 34 No. 1 Relative dating of the fossil hominids of Europe. K. P. Oakley. 1980. Pp. 1-63, 6 figs, 17 tables. £8.00
Vol. 34 No.2 Origin, evolution and systematics of the dwarf Acanthoceratid Protacanthoceras Spath, 1923

(Cretaceous Ammonoidea). C. W. Wright & W. J. Kennedy. 1980. Pp. 65-107, 61 figs. £6.25
Vol. 34 No.3 Ashgill Brachiopoda from the Glyn Ceiriog District, north Wales. N. Hiller. 1980. Pp. 109-216, 408
figs. £14.75

Vol. 34 No.4 Miscellanea: Upper Palaeozoic Athyrididae brachiopods—New British Cretaceous Epitoniidae—
Microproblematicum Prethocoprolithus—Glabellar structure of asaphid trilobites—New Lower Ordovician bivalve
family—Cretaceous brachiopods—Tupus diluculum sp. nov.—Revision of Plummerita. 1980. Pp. 217-297. £11.00

Vol. 35 No. 1 Lower Ordovician Brachiopoda from mid and south-west Wales. M. G. Lockley & A. Williams. 1981.

Pp. 1-78, 263 figs, 3 tables. £10.80
Vol. 35 No. 2 The fossil alga Girvanella Nicholson & Etheridge. H. M. C. Danielli. 1981. Pp. 79-107, 8 figs, 3 tables.
£4.20

Vol. 35 No.3 Centenary Miscellanea: Budleigh Salterton brachiopods—Oswald’s Turkish algae—J. A. Moy-
Thomas—Burials, bodies and beheadings—Nucleolites clunicularis—Phanerotinus cristatus—Fossil record of
teleosts—Neanderthal dating—Hippoporidra edax. 1981. Pp. 109-252. £20.00

Vol. 35 No.4 The English Upper Jurassic Plesiosauroidea (Reptilia) and a review of the phylogeny and classification
of the Plesiosauria. D. S. Brown. 1981. Pp. 253-347, 44 figs. £13.00

Vol. 36 No. 1 Middle Cambrian trilobites from the Sosink Formation, Derik—Mardin district, south-eastern Turkey.
W. T. Dean. 1982. Pp. 1-41, 68 figs. £5.80
Vol. 36 No.2 Miscellanea: Dinantian terebratulids—New microfossils—Neseuretus—Archaeocidaris whatleyensis—
Carboniferous dasyclad—Nanjinoporella—Toarcian bryozoans—Drybrook Sandstone plants—British fossil

bintoniellids—Uraloporella. 1982. Pp. 43-155. £19.80
Vol. 36 No.3 The Ordovician Graptolites of Spitsbergen. R. A. Cooper & R. A. Fortey. 1982. Pp. 157-302, 6 plates,
83 figs, 2 tables. £20.50

Vol. 36 No.4 Campanian and Maastrichtian sphenodiscid ammonites from southern Nigeria. P. M. P. Zaborski.
1982. Pp. 303-332, 36 figs. £4.00

Vol. 37 No. 1 Taxonomy of the arthrodire Phlyctaenius from the Lower or Middle Devonian of Campbellton, New

Brunswick, Canada. V. T. Young. 1983. Pp. 1-35, 18 figs. £5.00
Vol. 37 No. 2. Ailsacrinus gen. nov., an aberrant millericrinid from the Middle Jurassic of Britain. P. D. Taylor. 1983.
Pp. 37-77, 48 figs, 1 table. £5.90

Vol. 37 No.3. Miscellanea: Permian Glossopteris in Turkey—Wealden Theriosuchus—Wealden conifer—Permian
plants of Saudi Arabia—Carboniferous Edrioasteroidea—British cicadas—Dittonian cephalaspids. 1983. Pp. 79-171.
£13.50

Vol. 37 No.4 The relationships of the palaeoniscid fishes, a review based on new specimens of Mimia and Moy-
thomasia from the Upper Devonian of Western Australia. B. G. Gardiner. 1984. Pp. 173-428, 145 figs, 4 plates.

0 565 00967 2. 4 £39.00

Vol. 38 No. 1 New tertiary pycnodonts from the Tilemsi valley, Republic of Mali. A. E. Longbottom. 1984. Pp. 1-26,
29 figs, 3 tables. 0 565 07000 2. £3.90
Vol. 38 No. 2 Silicified brachiopods from the Viséan of County Fermanagh, Ireland. (III) Rhynchonellids, Spiriferids
and Terebratulids. C. H. C. Brunton. 1984. Pp. 27-130, 213 figs. 0 565 07001 0. £16.20
Vol. 38 No.3. The Llandovery Series of the Type Area. L. R. M. Cocks, N. H. Woodcock, R. B. Rickards, J. T-.
Temple & P. D. Lane. 1984. Pp. 131-182, 70 figs. 0 565 07004 5. £7.80
Vol. 38 No. 4 Lower Ordovician Brachiopoda from the Tourmakeady Limestone, Co. Mayo, Ireland. A. Williams &
G. B. Curry. 1985. Pp. 183-269, 214 figs. 0 565 07003 7. £14.50

Vol. 38 No. 5 Miscellanea: Productacean growth and shell shape—Jurassic alga Palaeosiphonium—Upper Ordovi-
cian brachiopods and trilobites—Lower Devonian Osteostraci from Podolia—Hipparion from Diavata—Preparation
and study of Singa skull—Carboniferous and Permian bryozoa—Lower Eocene trionychid—Montsech fossil insects.

1985. Pp. 271-412. 0 565 07004 5. £24.00
Vol. 39 No. 1 Upper Cretaceous ammonites from the Calabar region, south-east Nigeria. P. M. P. Zaborski. 1985.
Pp. 1-72, 66 figs. 0 565 07006 1. £11.00
Vol. 39 No. 2. Cenomanian and Turonian ammonites from the Novo Redondo area, Angola. M. K. Howarth. 1985.
Pp. 73-105. 33 figs. 0 565 07006 1. £5.60
Vol. 39 No.3 The systematics and palaeogeography of the Lower Jurassic insects of Dorset, England. P. E. S.
Whalley. 1985. Pp. 107-189, 87 figs, 2 tables. 0 565 07008 8. £14.00
Vol. 39 No.4 Mammals from the Bartonian (middle/late Eocene) of the Hampshire Basin, southern England. J. J.
Hooker. 1986. Pp. 191-478, 71 figs, 39 tables. 0 565 07009 6. £49.50
Vol. 40 No. 1 The Ordovician graptolites of the Shelve District, Shropshire. I. Strachan. 1986. Pp. 1-58, 38 figs. 0 565
07010 X, £9.00
Vol. 40 No. 2. The Cretaceous echinoid Boletechinus, with notes on the phylogeny of the Glyphocyphidae and Tem-
nopleuridae. D. N. Lewis. 1986. Pp. 59-90, 11 figs, 7 tables. 0 565 07011 8. £5.60
Vol. 40 No. 3 The trilobite fauna of the Raheen Formation (upper Caradoc), Co. Waterford, Ireland. A. W. Owen,
R. P. Tripp & S. F. Morris. 1986. Pp. 91-122, 88 figs. 0 565 07012 6. £5.60

Vol. 40 No.4 Miscellanea I: Lower Turonian cirripede—Indian coleoid Naefia—Cretaceous—Recent Craniidae—
Lectotypes of Girvan trilobites—Brachiopods from Provence—Lower Cretaceous cheilostomes. 1986. Pp. 125-222.
0 565 07013 4. £19.00
Vol. 40 No. 5 Miscellanea Il: New material of Kimmerosaurus—Edgehills Sandstone plants—Lithogeochemistry of
Mendip rocks—Specimens previously recorded as teuthids—Carboniferous lycopsid Anabathra—Meyenodendron,
new Alaskan lepidodendrid. 1986. Pp. 225-297. 0 565 07014 2. £13.00

Vol. 41 No. 1 The Downtonian ostracoderm Sclerodus Agassiz (Osteostraci: Tremataspididae). P. L. Forey. 1987. Pp.
1-30. 11 figs. 0 565 07015 0. £5.50
Vol. 41 No. 2. Lower Turonian (Cretaceous) ammonites from south-east Nigeria. P. M. P. Zaborski. 1987. Pp. 31-66.
46 figs. 0 565 07016 9. £6.50
Vol. 41 No.3 The Arenig Series in South Wales: Stratigraphy and Palaeontology. I. The Arenig Series in South
Wales. R. A. Fortey & R. M. Owens. II. Appendix. Acritarchs and Chitinozoa from the Arenig Series of South-west

Wales. S. G. Molyneux. 1987. Pp. 67-364. 289 figs. 0 565 07017 7. £59.00
Vol. 41 No. 4 Miocene geology and palaeontology of Ad Dabtiyah, Saudi Arabia. Compiled by P. J. Whybrow. 1987.
Pp. 365-457, 54 figs. 0 565 07019 3. £18.00
Vol. 42 Cenomanian and Lower Turonian echinoderms from Wilmington, south-east Devon. A. B. Smith, C. R. C.
Paul, A. S. Gale & S. K. Donovan. 1988. 244 pp, 80 figs, 50 pls. 0 565 07018 S. £46.50
Vol. 43. A Global Analysis of the Ordovician—Silurian boundary. Edited by L. R. M. Cocks & R. B. Rickards. 1988.
394 pp., figs. 0565 07020 7 £70.00
Vol. 44 No. 1 Miscellanea: Palaeocene wood from Mali—Chapelcorner fish bed—Heterotheca coprolites—Mesozoic
Neuroptera and Raphidioptera. 1988. Pp. 1-63. 0 565 07021 5 £12.00
Vol. 44 No. 2. Cenomanian brachiopods from the Lower Chalk of Britain and northern Europe. E. F. Owen. 1988.
0 565 07022 3 £20.70
Vol. 44 No.3. The ammonite zonal sequence and ammonite taxonomy in the Douvilleiceras mammillatum Superzone
(Lower Albian) in Europe. H. G. Owen. 1988. 0 565 07023 1 In press
Vol. 44 No. 4 Taxonomy and ecology of Cretaceous Cassiopidae. R. J. Cleevely & N. J. Morris. 1988. 0 565 07024 X
0 565 07024 X In preparation

Typeset by Santype International Limited, Salisbury, Wiltshire.
Printed in England by Oxford University Press.