GEOLOGICA CARPATHICA,  48, 3, BRATISLAVA,  JUNE 1997

145–161

DISTRIBUTION AND RECOGNITION OF PHASES IN APTIAN–TURONIAN

(CRETACEOUS) BRACHIOPOD DEVELOPMENT IN NW EUROPE

DANIÈLE GASPARD

Université de Paris-Sud, Département des Sciences de la Terre, Bât. 504,

91405 Orsay Cedex, France

(Manuscript received November 5, 1996; accepted in revised form March 18, 1997)

Abstract:

 Although represented during the Cretaceous (in three faunal provinces: Boreal, Jura, Tethyan), brachio-

pods have not commonly been considered as competitors of ammonites and foraminifers in the establishment of a
standard zonation scheme. Occasionally brachiopods are well represented in thin layers where other faunal groups are
absent or poorly represented. The paleontological evaluation (complementary morphological remarks with an illus-
tration and distribution of most species) is given here for the Aptian–Turonian period, and the correlative value of the
most important species is discussed. This analysis leads to a discussion–conclusion concerning: a) ecological consid-
erations (for example, the fact that some genera are preferentially found in typical facies), b) designation of strati-
graphic markers, c) location of the considered fauna elsewhere in the world and, d) a proposal for the recognition of
phases in brachiopod development in the present stage of knowledge, with remarks  concerning the distribution in
correlation with transgressive-regressive movements.

Key words: 

NW Europe, Aptian, Albian, Cenomanian, Turonian, recognition of phases, distribution, brachiopods.

Introduction

While well represented during the Cretaceous in the three
faunal provinces (Boreal, Jura and Tethyan), brachiopods
have not been considered as competitors of ammonites and
foraminifers in the establishment of a standard zonation. Be-
ing mainly anchored benthic organisms, they are highly con-
trolled environmentally and consequently their great varia-
tion may be responsible for their secondary zonal value. The
lack of recognition of micromorphic species which is due to
the small number of worldwide experts on the Cretaceous is
also a disadvantage.

It is not easy to correlate brachiopod occurrences with ammo-

nite zones, misleading, according to  Middlemiss (1981), but
this largely disregarded group has been collected in the past
without the ammonite fauna and without other faunal markers.
A better knowledge of successive faunas through time would
help to reach a true appreciation of ranges of species.

Each population frequently reflects a dimorphism, a point

particularly studied in the terebratulids (Sellithyridinae: Gas-
pard 1988). In addition, differences are observed between
populations of Phaseolina phaseolina (Lamarck) from the
Upper Cenomanian of Charentes, Sarthe, Vendée and Var
(France), the province of Beira (Portugal) and the Bohemian
Basin, a phenomenon known as polytypism. This phenome-
non is also observed, among other species, e.g. between pop-
ulations of Sellithyris cenomanensis (Gaspard) from Middle
Cenomanian strata from Sarthe, Indre and Charentes. This
variability in shape and size is related to local conditions. In
the case of rhynchonellids, with their decorative characters,
variability chiefly affects the ornamentation (mainly number
of costae) in addition to this external morphology.

Brachiopods are, however, occasionally well represented

in thin beds where major groups are absent. Thus they may

sometimes have local, regional or provincial correlative val-
ue. It does not appear necessary to redescribe entirely the
species, but for a better understanding of the discussion,
some complementary morphological remarks are presented
and placed in their systematic context. The stratigraphical
distribution of important genera and species  through Ap-
tian–Turonian strata is presented in Table 1. This will be re-
ferred to below in connection with the recognition of phases
in the brachiopod development and a preliminary zonation.

Paleontological evaluation: morphological

remarks and distribution

This section does not correspond to new systematic data,

but remarks on complementary morphological features are
sometimes necessary, as well as precise data on the distribu-
tion of taxa, prior to any discussion concerning their strati-
graphical value and recognition of phases of development.

Class Inarticulata Huxley 1869

Remarks:

  Inarticulates are not numerous in the Cretaceous,

few examples are quoted.

Superfamily Linguloidea

Family Lingulidae Menke 1828

Genus Lingula Brugière 1797

Remarks:

 Apart from Lingula subovalis Davidson briefly

recorded from the Upper Greensand near Warminster,
(Davidson 1852, pl. 1, Figs. 29, 30) and from Wiltshire
(Owen 1988, pl. 5), another species exists, Lingula raulini-
ana

  d’Orbigny 1847, which seems to be a synonym of L.

146                                                                                             GASPARD

truncata

  Sowerby described also by Davidson (1852, pl. 1,

Figs. 27, 28, 31). The shell of this species is thin, small,  ir-
regularly oval, flattened longitudinally, smooth with light
growth lines more numerous towards the commissures on
contrary of L. subovalis  with regular shape and few concen-
tric growth lines. It characterizes the Lower Greensand of
Atherfield (Isle of Wight) and the environs de Grandpré
(Ardennes).

Superfamily Cranioidea

Family Craniidae Menke 1828

Crania cenomanensis

 d’Orbigny  1847

Pl. II: Fig. 4

Remarks: 

The species has not been well studied in Europe.

It is known from the Upper Greensand (Cenomanian) of le
Mans (Sarthe). The description of the shell is based upon the
material of de Vibraye coll. (M.N.H.N., Paris, no. B. 16440)
because d’Orbigny coll., no. 6514, is non-existent now. From
this material, the shell appears irregular, approximately oval
transversally, the upper valve being somewhat conical. Mus-
cle scars, brachidium and a narrow limbus are well represented
on the internal surface of this valve (Pl. II: Fig. 4), while the
Lower valve is unknown.

Crania rhotomagensis

 d’Orbigny 1847

Contrary to the preceding species, C. rhotomagensis has a

longitudinally oval shell, with muscle fields closer to the
commissures and a wide limbus. It is known from Rouen in
d’Orbigny coll. (M.N.H.N., Paris no. 6515), but all speci-
mens are broken. Both species may, in fact, be representa-
tives of the genus Ancistocrania, according to the internal
characters, but more material is needed to prove this.

Class Articulata Huxley 1869

Family Rhynchonellidae Gray 1848

Subfamily Cyclothyridinae Makridin 1955

Genus Cyclothyris M’Coy 1844

Rhynchonella latissima

 J. de C. Sowerby 1829 is the type-

species of the genus.

Remarks: 

The species described by Owen (1962) occurs in

the Tropaeum subarcticum Subzone, Parahoplites nutfielden-
sis Zone of Berkshire, Cambridgeshire and Kent. Following
Barbulescu et al. (1975) it is also known to occur in the Ap-
tian of Baciuliu, Rasova (Rumania) and probably in the Jura
province.

Cyclothyris depressa

 (Sowerby 1825)

Remarks:

 D’Orbigny (1847) figured and described a com-

mon rhynchonellid species in synonymy with “R.” depressa
Sowerby, widely represented in France (M.H.N.H. coll., no.
5148). But comparisons of pl. 491 (Figs. 1–7) with the holo-
type figured by Owen (1962, pl. 4, Fig. 11) do not compare
favourably. The species is represented in P. nutfieldensis

Zone. “R.” lata specimens figured by d’Orbigny (M.N.H.N.,
no. 5149) seem to have more in common with “R.” depressa
Sowerby.

Cyclothyris antidichotoma

 (Buvignier 1843)

Remarks: 

The species, revised by Owen (1962), is repre-

sented in the Upper Aptian (P. nutfieldensis Zone) in En-
gland, with C. depressa (Upware, Cambridge, Brickhill,
Buckingamshire), as well in the Aptian of Ahaus (Pictet,
1872, pl. 199, Figs. 13–15). Barbulescu et al. (1975) quoted
the species from the Lower Aptian of Baciuliu Rasova (Ru-
mania). Though less common, the species is found in the
Lower Albian of Grandpré, Novion (Ardennes) (d’Orbigny
coll., no. 6014), Mont Saxonet, Goudinière, Savoie (France)
and Bedfordshire (England). The characteristic feature of the
shell of this species consists in grouping of fine costellae
(38–40) into strong costae from mid-length or third-anterior
towards the commissures.

Cyclothyris scaldisensis

 (d’Archiac 1846)

Remarks:

 The species is characterized by a shell broader

than long, biconvex, with a large circular foramen and a mas-
sive slightly incurved umbo. The ornamentation is composed
of numerous rounded costae. The species has been described
from the Lower Cenomanian (Tourtia of Tournai, Belgium).
It is recorded from the Lower Cenomanian of Normandy
(France), England and in the Munster Basin at Essen, Ger-
many. The material was observed in E.M. coll., Lyon.

Cyclothyris difformis

 (Valenciennes in Lamarck 1819)

Pl.  I: Fig. 15

Remarks:

 Apart from the type-series of Lamarck in Geneva,

the species has been collected from the Greensand (Lower
Cenomanian) near le Havre (Cap de la Hève). The specimens
are in M.N.H.N. coll., le Havre (no. 1250), (Pl. I: Fig. 15a–b).
Other specimens come from the Tourtia of Tournai (Lower
Cenomanian, Belgium) and le Mans (Lower and Middle Cen-
omanian, E.M. coll., Lyon), and some have been collected
from the Journeaux quarry (Valencay, Indre) in Middle Cen-
omanian. In addition to the previous localities, the species is
recorded from England, in the Lower Cenomanian deposits of
Essen, North Germany (Owen 1962) and probably in organo-
detritic limestones near Prague (Predboj, Bohemia).

The species name of Rh. difformis was controversialy em-

ployed. The species considered here has a variable shell shape,
being transverse or subtriangular in outline, with a few marked
growth lines. The width can attain 40 mm and the umbo is
sometimes short or massive, and more or less erect. Two dis-
tinct deltidial extensions in juveniles (Pl. II: Fig. 15), evoluate
to forme a typical circle around the foramen in adults. The
shell is ornamented by approximately 40 costae, fine near the
umbo, becoming coarser near the margins. The anterior asym-
metric margin is right or left-handed in adults.

The specimens from the Tourtia show certain resemblances with

C.

 scaldisensis, but the anterior region is more gibbous, having an

erect umbo and the anterior commissure asymmetrical.

DISTRIBUTION AND RECOGNITION IN APTIAN-TURONIAN BRACHIOPOD DEVELOPMENT                         147

It is necessary to separate all these Cenomanian specimens

from the Upper Cretaceous ones from Charentes, Landes,
Pyrenean region and S.E. of France, which also have an
asymmetrical commissure.

Cyclothyris compressa

 (Lamarck 1819)

Pl. I: Fig. 11

Remarks: 

Agreeing with the first presentation of Owen

(1962), I have observed this species in Lamarck coll. (pl. 19,
specimen A–C, Coulaines near le Mans, kept under the no.
46183), M.N.H.N. coll., no. 6495 pr.p., Paris; E.M. coll.,
Lyon. The species has been sometimes misidentified, its rep-
resentants have a winged shell, somewhat flat and transverse,
ornamented by 32 to 38 strong angular costae (7 in the fold)
with a characteristic thickening of the commissures (Pl. I:
Fig. 11b). The ventral, suberect, umbo is sharp dorsally. This
species characterizes a formation in the Sarthe region named:
“Les Sables du Perche”, (upper part of Middle Cenomanian–
lower part of Upper Cenomanian), Juignet (1974).

Cyclothyris lamarckiana

 (d’Orbigny 1847)

Pl.  I:  Fig. 17

Remarks: 

The species is also present in the Upper Cenoma-

nian, C. naviculare Zone “ Marnes à Ostrea biauriculata” of
the Sarthe region, but is slightly transverse, with a suberect
umbo. In comparison with C. difformis the ornamentation is
composed of approximately 33–35 well defined costae from
which, 6 are located in the fold (Pl. I: Fig. 17b) and the
asymmetry is not common. The foramen is circular and the
deltidial plates exposed (Pl. I: Fig. 17a). Representants of the
species have been observed in d’Orbigny coll., no. 6493,
Pèron coll., Morgan coll. (no. B. 16046), de Vibraye coll.,
no. B16052 (M.N.H.N., Paris).

Genus Burrirhynchia Owen 1962

Remarks: 

This name was created by Owen (1962) to distin-

guish “Rh”. leightonensis Walker (Bedfordshire, L. tardefur-
cata Zone) and “Rh.” cantabridgiensis Davidson (Aptian, P.
nutfieldensis Zone, Buckinghamshire),  from Cyclothyris.

The genus is also represented by B. gibbsiana (J. de C.

Sowerby 1826), redescribed by Owen (1956) (Lower Green-
sand of Folkestone) and B. sigma (Schloenbach 1867). As
distinct from the first three species, B. sigma (Lower Cenom-
anian) is more commonly found in Europe (North Germany,
Belgium, France, Poland, and also, Podolia). Popiel-Barczyk
(1977) has illustrated the species by serial transverse sections
of specimens from Poland (environs of Cracow, Czesto-
chowa, Iwanovice, Annopol). In addition to the pentagonal
outline, the shell is biconvex and the umbo suberect, but the
striking feature is the M-shaped anterior commissure charac-
terizing the species.

Subfamily Uncertain

Genus Grasirhynchia Owen 1968

Grasirhynchia grasiana

 (d’Orbigny 1847) is the type-spe-

cies of the genus. The lectotype selected by Owen (1968) is

figured from the d’Orbigny collection, no. 6497 of the cata-
logue (M.N.H.N., Paris). The species is recorded from
Warminster, Chute Farm, Isle of Wight (England), le Havre
and Pas de Calais (France).

Grasirhynchia martini

 (Mantell 1822)

Pl.  I: Fig. 16; Pl.  II: Fig. 16

Remarks: 

This species has been described by Owen (1968).

The shell is of small size, while G. grasiana is larger. Those
collected in 1978 and 1995 in Grey chalk of Blanc Nez cliffs,
between Escalles and Sangatte (Boulonnais) (upper part of
the Lower Cenomanian and Middle Cenomanian), particular-
ly in the bed above the 2nd O. mantelliana band, have the fol-
lowing dimensions (L: 6.0–8.4 mm; l: 5.8–7.4 mm; e: 3.7–
5.7 mm). The deltidial extensions around the foramen, although
not confined to the species, are well represented (Pl. II: Fig. 16).

Amédro (1993–1994, Fig. 3) indicated that the species is

found at the top of bed 17 and in bed 18 “Formation du Cran”
with  O. mantelliana Sowerby and Sciponoceras baculoides
(Mantell), in the same formation (set H’) with Micrabacia
coronula

 and other brachiopods including Modestella geinitzi

(Schloenbach) and Kingena concinna Owen.

The species is also recorded from the Aube region (Mont-

morency-Beaufort quarry) in a chalky marl with the previous
fauna (Amédro et al. 1994, Fig. 2).

Family Wellerellidae Likharev in Rzhonsnitskaya 1956

Subfamily Lacunosellinae Smirnova 1963

Genus Orbirhynchia Pettitt 1954

Orbirhynchia mantelliana

 (Sowerby 1826)

Pl.  I: Figs. 12–13

Remarks: 

This species is a very important stratigraphic

marker in the Cenomanian. Kennedy (1969) emphasized its
position in different locations of England, highlighting the
existence of an O. mantelliana band, stratigraphic marker,
occurring at the upper limit of the T. costatus Subzone, A.
rhotomagense Zone.

Kennedy (1969) quoted the species in Kent (coast): band 9

(mainly the top 30 cm) with Sciponoceras baculoides (Man-
tell),  A. rhotomagense (Brongniart) and Scaphites equalis
Sowerby, in band 10 (2.5 m) rich in specimens with Acantho-
ceras

; in Kent (inland), in the vicinity of Blue Bell Hill

Burham: band 4 (T. costatus fauna), in band 6 (O. mantelli-
ana

 band with Concinnithyris sp., Gemmarcula menardi

(Lamarck), Grasirhynchia grasiana (d’Orbigny) and Kinge-
na

 sp.); in Sussex, near Eastbourne; in Isle of Wight (O.

mantelliana

 band in Culver cliff and Compton Bay). The ex-

istence of this O. mantelliana band is summarized by
Kennedy (1969, p. 531). Pettitt (1949–54) quoted and figured
the species (p. 31, pl. 3, Figs. 10a–c) in Bedfordshire, Cam-
bridgeshire, Kent, Surrey, Sussex with an A. rhotomagense
Zone age.

The observations of Robaszynski et al. (1980) led to the

recognition of the existence of O. mantelliana in some local-
ities of Boulonnais, particularly at Escalles. Complementary
observations allow Amédro (1993, 1994) to define the exist-
ence of three O. mantelliana levels: one in the “Formation du

148                                                                                             GASPARD

Petit Blanc Nez”, set F, unit 10iv, containing also Monti-
clarella brevirostris

, another in the set G of the same forma-

tion, unit 15  associated with Rotalipora reicheli, the later in
set H’ of “Formation du Cran”, unit 18 (with Modestella
geinitzi

 (Schloenbach), Kingena concinna (Owen) and Gra-

sirhynchia martini

 (Mantell)) and mainly unit 19–19  with S.

baculoides

 (Mantell). The most important is the third level.

Field-trips in 1978 and 1995 gave again the opportunity of
collecting the species and accompanying fauna (Pl. I:
Figs. 12–13). In addition, material from France has been ob-
served in d’Orbigny coll. (M.N.H.N., Paris, no. 6911).

Comparisons have been made in Lower and Middle Cen-

omanian Chalks (Chalk Marl and Grey Chalk) between
Blanc Nez cliffs and at Folkestone (Kent), across the Straits
of Dover using Channel Tunnel boreholes correlations
(Amédro 1994, Fig. 5, p. 78). In the same manner accurate
lithological successions in the Paris Basin allow one to realize
the great spatial continuity of several marker levels in Middle
Cenomanian from Kent to Aube (Amédro et al. 1994).

Panow (1969, pl. CIX, Figs. 4a–c) quoted the species in

Korskiew and probably Sudol. Schloenbach (1867) men-
tioned the species from the Mantelliceras mantelli and Acan-
thoceras rhotomagense Zones, Northwestern Germany.

The Orbirhynchia boussensis-wiesti-cuvieri  group

Pl.  I: Figs.  2–4

The genus Orbirhynchia has been studied by Pettitt (1949-

50) in the context of the British chalk. Among other species
the author described: O. wiesti (Quenstedt), Cenomanian; O.
multicostata

 sp. nov., Turonian (Plenus marl); O. praedis-

pansa

 sp. nov. (Turonian, basal I. labiatus Zone); O. cuvieri

(d’Orbigny), basal  I. labiatus Zone (for a long time consid-
ered as a stratigraphic marker); some other new species from
I. labiatus Zone: O. extensa; O. compta; O. orbignyi; O. he-
berti

 (Terebratulina lata Zone); O. reedensis  Etheridge (Ho-

laster planus Zone). It seems that the species quoted after O.
cuvieri

 have no or a few representants elsewhere in Europe!

(e.g.  O. orbignyi reported from Poland by Popiel-Barczyk
1977). In this context, only O. wiesti (Quenstedt), O. cuvieri
(d’Orbigny) and O. boussensis Owen, a new species created
by Owen (1988), will be discussed.
Remarks: 

Comparisons from figured specimens of O. wiesti

and material from “Les Sables de Bousse” in the Sarthe De-
partment (sampled or observed in collections), as well as the
specimen of O. boussensis Owen shown here (Pl. I: Fig. 3)
allow to observe that the figured specimen of O. wiesti
(Owen 1988) is narrower than those from Bousse (Pl. I:
Fig. 4, but only one  specimen is shown!). O. cuvieri
d’Orbigny which characterizes the basal part of I. labiatus
Zone, is generally represented by specimens a little more
globular, similar to individuals from Rouen, (E.M. coll.,
Lyon) with the anterior commissure a little less uniplicate or
with an uniplication Lower than in specimens from Bousse.
The figuration of Pettitt from Surrey (1949–54, pl. III,
Fig. 12) and the lectotype chosen from d’Orbigny coll. (no.
6910a) do not show significant differences in the anterior
commissure. In addition, Juignet et al. (1973) quoted numer-
ous  O. wiesti in the type-section at Bousse (near the ceme-

tery). Peake & Hancock (1970, p. 306) do not clarify this dif-
ficult situation, quoting O. cuvieri and O. aff. wiesti in the I.
labiatus Zone of Norfolk. The number of costae does not al-
ways show a significant difference, leaving this situation as
it stands unless we consider a lineage in the process of time
with two main stages. However, it seems difficult to retain
two markers (O. boussensis and O. wiesti) for the upper Up-
per Cenomanian. Also, one should keep in mind the possibil-
ity of polymorphism and polytypism.

In addition, Middlemiss (1991) quoted the presence of rep-

resentants of the genus off Helgoland (North Sea).

Family Norellidae Ager 1959

Subfamily Monticlarellinae Childs 1959

Genus Monticlarella Wisniewska 1932

Monticlarella brevirostris

 (Roemer 1840)

Remarks:

 The species is known from Germany. Else, a few

specimens were sampled in 1995 from the first O. mantelli-
ana

 band, in glauconitic marly facies of Petit Blanc Nez

cliffs (Boulonnais) (L: 8.4–9.0 mm; l: 7.6–8.1 mm; e: 4.2–
4.9 mm). These specimens have a somewhat triangular shell,
with a ventral umbo slightly sharp and suberect and an oval
foramen. Bifurcating costae are observed at mid-length. A
widening is evident, close to the anterior limit of the shell.
All these characters allow to recognize this small species.

According to observations of Amédro (1993, 1994), 13

marker levels have been recognized from Blanc Nez to
Folkestone across the Straits of Dover. Amédro (1994,
Fig. 4) postulated a vertical extension of the species from
unit 3 to the top of unit 10 (with periods of abundance in
units 8 and 10) of “Formation du Petit Blanc Nez”.

Else Middlemiss (1991) highlighted the presence of the ge-

nus in North Sea (sea-floor off Helgoland).

Superfamily Terebratuloidea

Family Terebratulidae Gray 1840

Subfamily Sellithyridinae Muir-Wood 1965

Remarks: 

In this subfamily the genus Ovatathyris Owen

(1988) will not be discussed here without more precise data;
as well as the genus Boubeithyris (Cox & Midllemiss 1978).
B. boubei

 (d’Archiac) from the Tourtia of Tournai is the type-

species, B. diploplicata being evoked below as Praelongithy-
ris fecampi

 Gaspard.

Genus Sellithyris Middlemiss 1959

Remarks:

 The genus begins with S. sella (Sowerby) (type-

species) in the early Cretaceous, species several times men-
tioned notably in North Africa (Benest et al., 1996), mainly
in marly Hauterivian sediments and Lower Aptian in Spain,
France, England (Gaspard 1988, p. 88–107; Middlemiss
1968), Hungary (Dêtre 1968), (Pl. II: Fig. 13). The genus
ends with S. cenomanensis Gaspard, Middle Cenomanian
(type-locality: la Butte du Mans, Sarthe), in “Jalais” level
(Pl. I: Fig. 9). Both species are widely represented, although
intermediate species such as S. coxwellensis  (Middlemiss

DISTRIBUTION AND RECOGNITION IN APTIAN-TURONIAN BRACHIOPOD DEVELOPMENT                         149

1959),  S. longella (Leymerie 1869), and S. tornacensis
(d’Archiac 1846), (Pl. II: Fig. 1) are endemic: the first in the
Upper Aptian of England (Faringdon Sponge Gravel, Berk-
shire and Wiltshire), the second in the Upper Albian of Spain
and Pyrenean Greensand, the later in the Tourtia of Tournai,
Belgium (Lower Cenomanian). All species have been previ-
ously described (d’ Archiac 1848; Leymerie 1869; Peybernes
& Calzada 1977; Gaspard 1988) or analyzed by mutivariate
analysis (Gaspard & Mullon 1983) and thus do not require
complementary details in this context.

Genus Loriolithyris Middlemiss 1968

Remarks:

 For the period considered, the genus is represent-

ed by L. russillensis (Loriol) (type-species) ending in the be-
gining of the Aptian. This species, more characteristic of the
Jura and the Tethyan province (Gaspard 1988), is found dur-
ing the Aptian in southeast Morocco, eastern Spain (Mallada
1887), Alpes-Maritimes, Vercors and also Neuchâtel accord-
ing to Middlemiss (1981).

Genus Moutonithyris 1976

Type-species: M. moutoniana (d’Orbigny 1847)

Pl. II: Fig. 9

Remarks:

 The species is known from the Berriasian (mainly

South of France) but its distribution until the Aptian of Ger-
many, Westphalen (Schloenbach 1860) and probably Sardinia
(Dieni et al. 1973) is interesting in this context. The genus
becomes extinct with M. dutempleana (d’Orbigny), Upper
Albian-basal Cenomanian (Gault-Upper Greensand), (Pl. II:
Fig. 2). While ubiquitous, this species is well represented in
the Boulonnais (France), as well in  England (Gaspard 1976,
1985, 1988, p. 197–201) apart the Tethyan realm.

Representatives of both M. moutoniana and dutempleana

are quoted from North Sea (Middlemiss 1991).

The species M. obtusa (Sowerby 1823) seems to be possi-

bly confined to Cambridge Greensand.

Genus Phaseolina Gaspard 1985

Remarks:

 The genus presumably terminates the Sellithyridi-

nae. At the moment, obviously, only one species is known: P.
phaseolina

 (Lamarck).

Phaseolina phaseolina 

(Lamarck 1819)

Pl. I: Fig. 5

The species, widely described in several populations (Gas-

pard 1983, 1985, 1988) is representative of the Upper Cen-
omanian (type-locality, Colline de la Goupillerie, Sarthe in
“Sables ˆ Catopygus obtusus”, Pl. I: Fig. 5), sometimes in
company of Orbirhynchia  group boussensis-wiesti-cuvieri
(cf. “Sables de Bousse” lateral extension of the first forma-
tion), in other locations with Ostrea biauriculata (Maine &
Loire) or Calycoceras, and in extreme position with Gem-
marcula carentonensis

 (d’Orbigny) (Pl. I: Fig. 6) and Arca

(Charente Maritime) (Gaspard 1985, 1988). It is also quoted
in the Bohemian Basin (Nekvasilova 1973) and observed by

the present author from the Late Cenomanian in the collec-
tions of the Geological Institute of Freiberg University of
Mining and Technology (Germany).

Genus Praelongithyris Middlemiss 1959

Remarks:

 The genus extends from the Aptian to the Cenom-

anian, with P. praelongiforma Middlemiss (1959) from the
Upper Aptian (abundant at Upware, less present at Brickhill
(?), and Faringdon) and P. lankesteri Walker (1868) (both
possibly endemic in England), through the Lower Albian : P.
rogeri

 (Gaspard 1974, Holotype selected from d’Orbigny

collection, M.N.H.N., Paris, no. 6017A), in Leymeriella tar-
defurcata Zone (les Ardennes, France) (Pl. II: Fig. 6), (Gas-
pard 1988, p. 210–218), to the Lower Cenomanian (H. carci-
tanensis–M. saxbii Zones) with P. fecampi Gaspard in
Nièvre, Normandy, (Pl. II: Fig. 5), probably Tourtia of Tour-
nai (I.R.S.N, Brussels, no. IG 10511/20, 21) and Southern
England (Pl. II: Fig. 10; Gaspard 1985, 1988). This last spe-
cies was described under the name of Boubeithyris diplopli-
cata

 by Owen (1988, p. 112, pl. 8).

Subfamily Rectithyridinae Muir-Wood 1965

Genus Rhombothyris Middlemiss 1959

Remarks: 

The species Rhombothyris extensa (Meyer 1864)

is known from the Upper Aptian of Upware, Brickhill, Bar-
gate beds (P. nutfieldensis Zone, West Surrey) and glauconitic
sands of Shanklin (Isle of Wight). R. microtrema (Walker
1868) was found in the two first localities, R. meyeri (Walker
1868) at Upware and R. conica Middlemiss (1959) at Brick-
hill. All these species seem to be confined in England.

Genus Cyrtothyris Middlemiss 1959

Remarks: 

As for the previous species, representatives are

known in the Aptian of England (C. uniplicata (Walker 1870),
C. cantabridgiensis 

(Walker 1870), C. seeleyi (Walker 1870),

C. dallasi

 (Walker 1867), and also from Germany (Berklin-

gen), Switzerland (Ste. Croix), C. cyrta (Walker 1868).

Genus Rectithyris Sahni 1929

Remarks: 

This genus may have been evolved from Cyr-

tothyris 

Middlemiss 1959  (Middlemiss 1984). It is repre-

sented by R. viquesneli (d’Archiac 1846), Upper Albian, R.
depressa

 (Lamarck 1819), Lower Cenomanian (Tourtia of

Tournai, Belgium) and has a restricted distribution (Pl. II:
Fig. 3). This species was observed in the following collec-
tions (I.R.S.N., Brussels; M.N.H. N., Paris; E.M., Lyon.).

Genus Tropeothyris Smirnova 1972

Remarks: 

This genus not well represented, will not be dis-

cussed without more concrete data (cf. Owen 1988).

Genus Concinnithyris Sahni 1929

Remarks:

 Few papers concern the genus Concinnithyris pre-

viously placed in Gibbithyridinae. One of the last papers

150                                                                                             GASPARD

(Middlemiss, 1991 p. 225) argued to place this genus in the
subfamily Rectithyridinae; more investigations are needed to
adopt definitively this position. The genus is not confirmed
to be a transition from Moutonithyris (Middlemiss 1984,
1991), and is widely represented particularly in the Cenoma-
nian and Turonian of Normandy, South England, Germany?,
with C. albensis (Leymerie), C. obesa (Sowerby), C. subun-
data

 (Sowerby) and C. protobesa Sahni.

Concinnithyris obesa

 (Sowerby 1823)

Pl.  I: Fig. 18

Remarks:

 This is an example for which the interpretation of

the species is problematic because of the great variability of
the global morphology of the shell. The propensity of Sahni
(1929) to create many species around C. obesa does not fa-
cilitate discussions. The type-specimen probably does not re-
flect the median aspect of the species but the variants are so
numerous that one can find medium-sized specimens, gib-
bous, but narrow with nearly parallel flanks or lateral com-
misures and sometimes a gentle beak angle (d’Orbigny
1847–51, pl. 513, Fig. 4).

These large or medium specimens, round gibbous or nar-

row, in collections, come from “La Craie de Rouen”, Côté
Ste Catherine (Normandy), (d’Orbigny coll. no. 6912) which
is now recognized as Middle Cenomanian (Juignet 1974).
Some large specimens come from Neuvy (Yonne, E.M. coll.)
and St. Sauveur (Yonne; d’Orbigny coll., M.N.H.N., Paris).
Owen (1988) also recognized this great variability.

To conclude, the object is to obtain a great number of

specimens for multivariate analysis, from each locality, to
determine (once and for all), the presence or not of one or a
few other species.

Concinnithyris protobesa

 Sahni 1929

Remarks:

 This species has external morphological affinities

with the narrow figured specimen from “La Craie de Rouen”
in d’Orbigny, as well as with the specimen in Owen (1988,
pl. 14) and with some specimens of C. obesa from the E.M.
coll., Lyon. In other respects the specimen figured by David-
son (1852–54) has little to do with the type-species C. proto-
besa

 present in the Lower Turonian of England and Nor-

mandy, which has a shell characterized by a well curved beak
and appressed to the dorsal umbo preventing the symphytium
to be exposed. The foramen is large, circular or ovate and
sometimes slightly labiate. The brachial valve is less convex
than the pedicle one. The anterior margin is in intermediate
expression between retimarginate and uniplicate.

Concinnithyris  subundata

 (Sowerby 1813)

Pl.  I: Fig. 19

Remarks:

 This species seems at first sight to be more easy

to distinguish from C. obesa, because of the less biplicate
and gibbous anterior commissural region while the shell
shape is subpentagonal or subcircular. The species is repre-
sented in the Lower Chalk in England (Kennedy 1969; Peake
& Hancock 1970) and France (particularly in Normandy and
Boulonnais where juveniles specimens have been sampled).

Plate I: Fig. 1

 — Terebratulina lata Etheridge (1881), Holotype,

marker of the Middle Turonian (T. lata Zone),  (British Geological
Survey coll.) BSG GSM, no. 38693, 

×

4. Fig. 2a–b — One speci-

men of the genus Orbirhynchia  from “Sables de Bousse”, Bousse,
Sarthe (France),  pers. coll., (a) dorsal view, (b) anterior view,

×

1.2.  Fig. 3 — O. boussensis Owen (1988), from “Sables de

Bousse”, Bousse, Sarthe (France), (anterior commissure), (British
Museum): B.M (N.H.) coll. no. BB 82131, 

×

1.2. Fig. 4a–b — Orbi-

rhynchia wiesti 

(Quenstedt), Upper Cenomanian, Bed B, Little

Beach Beer specimen figured by Owen (1988) (a) dorsal view, (b)
antrerior view. Coll. B.M.(N.H.) coll. no. BB 7126, 

×

1.2 (for com-

parison with Pl. I. 2,3). Fig. 5 — Phaseolina phaseolina (Lama-
rck), “Sables à Catopygus obtusus”, Upper Cenomanian of
Mezieres-sous-Ballon, Sarthe. Pers. coll., 

×

1.2. Fig. 6 — Gem-

marcula carentonensis

 (d’Orbigny), “Horizon A”, upper Cenoma-

nian, Charente Maritime (France). (Collection of Ecole des Mines,
location: Faculté des Sciences de Lyon), E.M. coll. FSL no. 20301,

×

1.2. Fig. 7 — Kingena elegans  Owen (1970), Holotype, Turoni-

an (Holaster planus Zone). B.M. (N.H.) coll.  no. B55241, 

×

1.2.

Fig. 8

 — Dereta pectita (Sowerby), from  Upper Greensand (Cen-

omanian), near le Havre (France), E.M. coll., FSL no. 20302, 

×

1.2.

Fig. 9

 — Sellithyris cenomanensis (Gaspard), from “Jalais” level

(Middle Cenomanian), upper part of “Sables et Grés du Mans”,
Sarthe (France). E.M. coll. FSL no. 20303, 

×

1.2. Fig. 10 — Gem-

marcula menardi

 (Lamarck), same level and location as Pl. I. 9.

E.M. coll., FSL no. 20304, 

×

1.5. Fig. 11a–b — Cyclothyris com-

pressa

 (Lamarck), characteristic of “Sables du Perche”, Sarthe,

France.  (a) dorsal view, (b) anterior view. E.M. coll., FSL no.
20305, 

×

1. Fig. 12 — Orbirhynchia mantelliana (Sowerby), Low-

er Chalk, Middle Cenomanian (A. rhotomagense Zone, T. costatus
assemblage, Eastbourne (Sussex), England). Kennedy coll., B.M.
(N.H.), no. BF 50, 

×

1.2. Fig. 13 —  O. mantelliana (Sowerby), A.

rhotomagense

 Zone, 2nd. O. mantelliana band,  Boulonnais (be-

tween Escalles and Grand Blanc Nez). Pers. coll., 

×

1.4. Fig. 14 —

 Modestella geinitzi (Schloenbach), above the 2nd O. mantelliana
band (mid. Cenomanian), Blanc Nez Cliffs (Boulonnais). Pers. coll.,

×

1.5. Fig. 15a–b — Cyclothyris difformis (Lamarck), Lower Cen-

omanian, Normandy, (a) dorsal view, (b) anterior view. Mus.
Havre coll., no.1250, 

×

1.2. Fig. 16 — Grasirhynchia martini

(Mantell), same level and location as Pl. I. 14. Pers coll., 

×

1.75.

Fig. 17

 — Cyclothyris lamarckiana (d’Orbigny), le Mans (Sarthe),

“Marnes  à  O. biauriculata”. (a) dorsal view, (b) anterior view.
E.M. coll., FSL no. 20307, 

×

1. Fig. 18 — Concinnithyris obesa

(Sowerby), from “La Craie de Rouen”, Côte Ste Catherine, Nor-
mandy. E.M. coll.,  FSL no. 20306, 

×

1. Fig. 19 — Concinnithyris

subundata

 (Sowerby), Lower Chalk, Middle Cenomanian (A. rho-

tomagense Zone, O. mantelliana band, Abbotscliff, Folkestone
(Kent, England)). Kennedy coll. B.M. (N.H.), no. BB 85781, 

×

1.

Fig. 20a–b

 — Gibbithyris semiglobosa (Sowerby), Turonian,

Villevallier (Yonne, France: (a) dorsal view, 

×

1.2, (b) anterior

view of another specimen, Pers. coll. 

×

1.

Representatives of the genus have been quoted by Middle-

miss (1991) in the North Sea and by Kennedy (1969) in the
Middle Cenomanian, but often without determination in this
last case. More often it is C. albensis (Leymerie) (Kennedy
coll. in Br. Mus. Nat. Hist., no. B. 85783–4), or C. subundata
(no. BF 48–49, BB 85781–2) from Kent, Sussex, Dorset (cf.
A. rhotomagense Zone, Orbirhynchia mantelliana band).
Kennedy (1969) highlighted the occurrence and particular
abundance of Concinnithyris and sometimes Gibbithyris at
the boundary of T. acutus with A. jukesbrownei assemblages.

→

PLATE I                                                                                                 151

152                                                                                             GASPARD

Subfamily Gibbithyridinae Muir-Wood 1965

Genus Gibbithyris  Sahni 1925

Two species are of interest for the periods concerned: G.

semiglobosa

 (Sowerby) and G. subrotunda (Sowerby).

Gibbithyris semiglobosa

 (Sowerby 1813)

Pl. I: Fig. 20a–b

Remarks:

 The species is quoted in Turonian by Sahni

(1929), in British Chalks, in “La Craie de Meudon” (Paris
Basin) by d’Orbigny (no. 7676 in his catalogue) and in the
White Chalk from Fécamp (France). Specimens have been
collected in the Turonian of Villevallier (Yonne, France) with
a round shape biconvex, curved beak, minute circular fora-
men and a very gentle biplicate anterior commissure (Pl. I:
Fig. 20b). Microstructure analysis of the shell reveal a pris-
matic tertiary layer.

Gibbithyris subrotunda

 (Sowerby 1813) seems to have the

same vertical distribution as G. semiglobosa, and is better
known from the Turonian. The species is characterized by a
more appressed beak against the dorsal umbo and a greater
breadth which involves a pronounced break with lateral com-
missures on contrary of G. semiglobosa.

Subfamily  Capillithyridinae Cooper 1983

Genus Capillithyris Katz 1974

Remarks: 

Erroneously identified as Platythyris (Popiel-

Barczyk 1972; Middlemiss 1978; Bilinkevich & Popiel-
Barczyk 1979). The genus is represented by several species,
essentially from the Cenomanian: C. capillata, C. squamosa,
C

.  disparilis. Representatives of the genus are quoted also

from the North Sea.

Capillithyris capillata

 (d’Archiac 1846)

Pl. II: Fig. 8

Remarks: 

Found in the Tourtia of Tournai (Belgium), basal

Cenomanian, the species has a round shell, a large foramen
and a slightly convex brachial valve. Strong growth lines are
characteristic of the genus, and fine radiating lines give a
regular wavy ornamentation  at mid-length.

Capillithyris squamosa

 (Mantell 1822)

Pl.  II: Fig. 17

Remarks:

 Shells of medium size have been sampled in le

Boulonnais in bed 17 (Middle Cenomanian) with particular
growth-lines which give to this shell a regular step-like as-
pect. Fine radiating lines, better observed near the commis-
sures, complete the ornamentation (Pl. II: Fig. 17).

Capillithyris disparilis

 (d’Orbigny 1847)

Pl. II: Fig.  7

Remarks: 

This species sampled from Rouen presents an

oval shell, elongated, gibbous in the half-posterior part, with

Plate II: Fig. 1

 — Sellithyris tornacensis (d’Archiac), Tourtia de

Tournai, Montignies-sur-roc (Belgium). E.M. coll., FSL no.
20308, 

×

1. Fig. 2 — Moutonithyris dutempleana (d’Orbigny),

Barrington pit Cambridge, (Upper Greensand). Pers. coll. 

×

1.

Fig. 3

 — Rectithyris depressa (Lamarck), Tourtia of Tournai,

Montignies-sur-roc, Belgium. E.M., coll., FSL no. 20309, 

×

1.

Fig. 4

 — Crania cenomanensis d’Orbigny, “Jalais” level, le Mans

(Sarthe). de Vibraye coll. M.N.H. N., Paris, no. B.16440, 

×

1.5.

Fig. 5

 — Praelongithyris fecampi Gaspard, H. carcitanensis and

M. saxbii 

Zones, Val aux Clercs near Fécamp  (type-locality,

France). Mus. Rouen coll., no.  60473a, 

×

1. Fig. 6 — Praelon-

githyris rogeri

 Gaspard, Leymeriella tardefurcata Zone (Lower

Albian, Grandpré, Ardennes, (France). E.M. coll., FSL no.
291 014, 

×

1. Fig. 7 — Capillithyris disparilis (d’Orbigny), Rouen

(France). E.M. coll., FSL no. 20310, 

×

1.3. Fig. 8 — Capillithyris

capillata

 (d’Archiac), Tourtia of Tournai, Montignies-sur-roc

(Belgium), I.R.S.N. coll. (Institut Royal des Sciences Naturelles
de Belgique), 

×

1. Fig. 9 — Moutonithyris moutoniana (d’Orbigny),

Neocomian Chartreuse, 

×

1. Fig. 10 — Praelongithyris fecampi

Gaspard, Upper Greensand, Warminster, 

×

1.5. Fig. 11 — Terebratu-

lina protostriatula

 Owen, Lower Cenomanian (glauconitic marl),

from St. Catherine’s cliff above Rocken End (Isle of Wight). Evans’
Coll., 

×

1.5. Fig. 12 — Terebratulina martiniana d’Orbigny, Albian,

Martigues (Bouches du Rhône, France). d’Orbigny coll., M.N.H.N.,
Paris, no. catalogue 6018 (B. 14746). 

×

1.5. Fig. 13 — Sellithyris

sella

 (Sowerby), Lower Aptian of I. of Wight. Pers. coll., 

×

1.5.

Fig. 14

 — Argyrotheca pectunculus Quenstedt from the Upper

Greensand near Neufchatel (Boulonnais). E.M. coll., FSL no.
20311, 

×

8. Fig. 15 — Cyclothyris difformis  (Lamarck), posterior

part of a juvenile shell from the Tourtia of Tournai (Belgium) show-
ing the extensions of the deltidial plates around the foramen, 

×

18.

Fig. 16

 — Grasirhynchia martini (Mantell), posterior part of a shell

from bed 17 in “Formation du Cran” (Boulonnais), showing the ex-
tension of the deltidial plates or auriculations, 

×

30. Fig. 17 —

 Capillithyris squamosa (Mantell), upper part of bed 17 “Formation
du Cran” (Boulonnais), exhibiting regular growth lines and a fine
radiation more visible near the commissures, 

×

5.

strong growth lines and a light ornamentation in zigzag to-
wards the margins. The material was observed in d’Orbigny
coll., no. 6503; de Vibraye coll., no. B 16056 and Péron coll.,
no. S 07523 (M.N.H.N., Paris).

Superfamily Cancellothyridoidea

Family Cancellothyrididae Thomson 1926

Subfamily Cancellothyridinae Thompson 1926

Genus Terebratulina d’Orbigny 1847

Terebratulina martiniana  

d’Orbigny 1847

Pl. II: Fig. 12

Remarks:

 The species, observed in d’Orbigny coll., no. cat-

alogue 6018 (M.N.H.N., Paris, general coll., no. B 14746) is
defined as a little triangular shell, with a brachial valve less
convex than the pedicle one, ornamented by fine costellae ra-
diating from mid-length. The species is found in the Albian
of Martigues (Gueule-d’Enfer, Bouches du Rhône, S.
France) with Gryphea (Pl. II: Fig. 12) and near Villars de
Lans (Isère, France). Dieni & Massari (1965) recorded the
species with Acanthoplites nolani (Upper Aptian) from
Orosei (Sardinia), while Dieni et al. (1973) recorded it from
mid-Aptian to Upper Albian from France, England, Germany

→

PLATE II                                                                                              153

154                                                                                             GASPARD

and Sardinia. Middlemiss (1991) recorded the species off
Helgoland (North Sea).

Terebratulina protostriatula

 Owen 1988

Pl.  II: Fig. 11

Remarks:

 This species has been created to differentiate the

Cenomanian forms from T. striatula  (Mantell 1852), Upper
Chalk. The species occurs in the Lower Cenomanian of En-
gland, Boulonnais and Normandy (France). Material of the
British Geological Survey (BGS) has been observed, is com-
posed of elongate shells ornamented with fine costae.

In addition to T. martiniana and T. protostriatula, two spe-

cies of Terebratulina: T. triangularis Etheridge (1881), (Up-
per Greensand, Cambridge) and T. nodulosa Etheridge
(1881), (Upper Albian to Lower Cenomanian and (?) Middle
Cenomanian appear to be present in the material from Nor-
mandy (B. Ferré, Ph. D., University of Paris 6, Jussieu
1995).

Terebratulina lata

 Etheridge 1881

Pl. I: Fig. 1

This species (ex variety of T. gracilis ) has been created to

prevent any confusion with the species Terebratulina gracilis
Schlotheim previously described from the Senonian. T. lata
is characterized by a somewhat plano-convex shell, round in
outline, approximately 6 mm in length, with well observed
deltidial plates and strong radiating nodulous costae dividing
several times. This species, a stratigraphic marker in British
Chalks (Middle Turonian), is considered in parallel with Col-
lignoniceras woolgari

  (Wright & Wright) in international

Ammonite Zones (Rawson et al. 1978). T. lata is observed
particularly in North and Eastern part of France (d’Orbigny
coll., no. 6914) and in England (Middle Chalk of Babraham
road near Cambridge, at White Cliff near Seaton (Devon),
Swanage Bay (Dorset), Buckinghamshire). The type-materi-
al is kept in the British Geological Survey, Nottingham, un-
der the following registered numbers: no. BSG GSM 38693
(Holotype, Pl. I: Fig. 1), BSG GSM 29422–29433, BSG GSM
119253–119257.

Superfamily Zeillerioidea

Family Zeilleriidae Allan 1940

Subfamily Cheirothyridinae Baker

Genus Modestella Owen in Casey 1961

Modestella geinitzi 

(Schloenbach 1866)

Pl. I: Fig. 14

Remarks:

 After the revision by Owen (1988), I have sam-

pled, in 1995, some new specimens of the species in Blanc
Nez cliffs (Boulonnais). The shell is small, gibbous (L: 11–
8.3; l: 10.5–7.4; e: 7.3–4.6 mm) approximately round pentago-
nal, with a large circular foramen. Some pronounced growth
lines are situated in the third-anterior part of the shell. By its
abundance the species is considered as one of the biomarkers
in the Lower and Middle Cenomanian chalks across the
Straits of Dover (Amédro 1993, 1994) and localised in “For-

mation du Cran”: units 17 and 18, where it occurs with Kin-
gena concinna

 (Owen), Grasirhynchia martini (Mantell)

and, some Orbirhynchia mantelliana  (Sowerby) at the top of
unit 18.

Apart from North of France and England, the species is

quoted from the Greensand of Essen, North Germany and the
Cenomanian of the environs of Annopol (Poland).

Superfamily Megathyridoidea MacKinnon & Smirnova 1995

Family Megathyrididae Dall 1870

Genus Argyrotheca Dall 1900

Remarks:

 In addition to “Argyrotheca” megatrema (Sowerby

1836) described by Owen (1988) from Cambridge and from
Warminster, another species, probably of Megathyridoidea,
was observed in the E.M. coll., from the Upper Greensand of
Boulonnais near Neufchâtel. This species considered as “T”
pectunculus 

Quenstedt 1885 is very few ornamented (5–6

costae with few intercalations, Pl. II: Fig. 14), with less cos-
tae than in A. megatrema. No papillae are observed, but a
lamellar aspect enhances the ornamentation. The few speci-
mens in collections prevent us to practice transverse serial
sections on the material, so no observations have been real-
ized on the brachidium.

Superfamily Terebratelloidea

Family Dallinidae Beecher 1893

Subfamily Gemmarculinae Elliott 1947

Genus Gemmarcula Elliott 1947

Gemmarcula aurea

 Elliott 1947

The type-species illustrated by Owen (1977, p. 211, pl. 1)

is an endemic species from the  Aptian (Lower Greensand) of
Britain, confined to the Sponge Gravel of Faringdon. The
species has some resemblance to  G. crassicosta (Leymerie).

Gemmarcula crassicosta

 (Leymerie 1869)

Remarks: 

The plano-convex shell of this species from Vim-

port (France) is recognized in the type-level: Lower Clan-
sayesian (Simplorbitolina manasi Zone) by Peybernes &
Calzada (1975).

Gemmarcula menardi

 (Lamarck 1819)

Pl. I: Fig. 10

Remarks:

 The type-series is in the Lamarck collection (M.

H. N., Geneva) and is figured  in the catalogue of Favre
(1918, pl. 16, Fig. 93 (A-D)-99), collected from Coulaines
near le Mans (Middle Cenomanian). Owen (1977, p. 215) se-
lected the lectotype as specimen A.

In d’Orbigny collection (M.N.H.N., Paris, no. 6505), and

in Deshayes coll. (E.M.), the species is well represented and
following the aspect of the matrix probably comes from the
“Jalais” level in the Sarthe region, France (Middle Cenoma-
nian), (Pl. I: Fig. 10).

In his thesis, Juignet (1974) referred to the presence of G.

menardi

 in “Sables et Grès de la Trugalle et Lamnay”

DISTRIBUTION AND RECOGNITION IN APTIAN-TURONIAN BRACHIOPOD DEVELOPMENT                         155

(Sarthe); at Savigné-l’Evêque with Trigonia sulcataria
Lamarck under the hard-ground Longeville 2, with O. colum-
ba 

var minor  Deshayes (upper part of hard-ground Longev-

ille 1), at Parancé in Loyer quarry, at Lamnay in Louvre
quarry, at Théligny in la Rouaudière quarry in “Sables et
Grès de la Butte du Mans”; in the Lower part of “Sables du
Perche” (Sarthe) in cavities of the “Jalais” level, A. jukes-
brownei Zone. In any case, this species characterizes the up-
per part of the Middle Cenomanian.

In addition to the topotypes, G. menardi is found in the Mid-

dle Cenomanian of Valencay (Indre, Alcaydé 1966), in sandy
glauconitic facies, at Langé in the same beds as Sellithyris
cenomanensis

 Gaspard as in Sarthe, also with Cyclothyris,

Trigonia sulcataria

 Lamarck, T. crenulata Lamarck, Neithea

quinquecostata

 d’Orbigny, and Catopygus columbarius

d’Archiac. Cephalopods collected in these beds are Turrilites
costatus

 Lamarck, Euomphaloceras sussexiense Sharpe,

Acompsoceras rothomagense

 Brongniart, all from the Middle

Cenomanian, and showing similarities with the Sarthe region.
Unfortunately, the species is not found in other localities else-
where in France where S. cenomanensis Gaspard is present.

According to Kennedy (1969), G. menardi is found in En-

gland, in the Kent (coast): band 6 in the A. rothomagense Zone
with Concinnithyris albensis (Leymerie). The species was also
collected in the region of Burham (Kent) in the Orbirynchia
mantelliana

 band (cf. tabl. IV.) as well as near Holborough Ce-

ment Works. The sections in Isle of Wight (Culver Cliff and
Compton Bay) contain the species.

Specimens of G. menardi have been recorded from the

North of Germany. Figs. 6a–c, pl.1 of Popiel-Barczyk (1972)
correspond to the species (Poland, Cracow region), but in
other respects the citation of the species in Panow (1969)
seems to be a misidentification.

Gemmarcula carentonensis

  (d’Orbigny 1847)

Pl.  I: Fig.  6

Remarks: 

This  species is characterized by a large shell, some-

what broader than long, an antiplicate anterior commissure, a
median sulcus in the pedicle valve originating from the umbo
and two faint folds, as well a suberect umbo with distinct beak
ridges, a small circular submesothyrid foramen, and a distinct
area. The ornamentation consists of numerous radiating costae
(40) with bifurcations, some more marginal than others.

D’Orbigny assigned the species to the Ammonites mantelli

(?) Zone at Port des Barques (Charentes, France) (d’Orbigny
coll., M.N.H.N., no. 6506) and Eoux (Basses Alpes, France).
Publications of Kennedy & Juignet (1973) and Juignet et al.
(1973) define exactly the situation, localising “La Craie à
Terebratella carentonensis

”: as Horizon A in parallel with

the nodulous Chalk with Sciponoceras gracile Shumard,
Kanabiceras septemseriatum 

 (Cragin) and Inoceramus gr.

pictus

 in Upper Normandy, as inferior part of Melbourn Rock

with  S. gracile in SE England and Faune remaniée with
“Neocardioceras Pebble Bed” at the basal part of Middle
Chalk in the Devon coast.

Subfamily Uncertain

Genus Arenaciarcula Elliott 1959

Remarks:

 Apart from A. fittoni (Meyer 1864) typical of the

Upper Aptian of S England, another species is recorded from
the Cenomanian of Europe it is:  Arenaciarcula beaumonti
(d’Archiac 1846).

Recognized in the Tourtia of Tournai, Belgium, by d’Archiac,

the species has been revised by Popiel-Barczyk (1972) from An-
nopol (Poland), but was distinguished from Oblongarcula by
Owen (1977). Geographically widespread, the species is recog-
nized also in the Essen Greensand of North Germany, France
(Middle Cenomanian of Indre for example), Denmark, and in
the Cenomanian limestone of Devon and Russia.

Family Terebratellidae King 1850

Subfamily Trigonoseminae Elliott 1965

Genus Dereta Elliott 1959

Dereta pectita 

(Sowerby 1816)

Pl.  I: Fig. 8

The shell of medium size is smaller than either Gemmarcula

carentonensis

 or G. menardi, with which it migh have been

confused, and slightly biconvex. The beak is suberect with a
circular foramen, larger than in G. carentonensis, but less than
in G. menardi, and a straight hinge-line is observed. Numerous
costae (around 40), thinner than in G. menardi and G. caren-
tonensis

, with bifurcation at mid-length of the shell, constitute

the ornamentation. The sulcation and folds of the anterior
commisure are light. The holotype comes from the Upper
Greensand of Horningsham (Wiltshire). Apart from the type
locality, the species is found near Warminster, quoted also
from the Mantelliceras mantelli Zone (?) (Kennedy 1969) at
Compton Bay (Isle of Wight) in glauconitic marl Dorset.

The specimens observed in the Kennedy collection (Br.

Mus.), in d’Orbigny coll. (no. 6507) and in Deshayes coll.
(E.M., Lyon) are smaller than the type, and come from le
Havre and other localities in the glauconitic marl on Nor-
mandy coast (France).

Subfamily Uncertain

Genus Terebrirostra d’Orbigny 1847

Terebrirostra arduennensis 

d’Orbigny 1847

Remarks:

 The long pedicle umbo gives the shell an accen-

tuated elongate aspect characterized by the aptitude of spec-
imens of the genus to colonize soft substrates. The shell is
covered by dichotomous costae. The species was described
from the Lower Albian of Grandpré (Ardennes, France) by
d’Orbigny (1847, pl. 519, Figs. 6–10) and by Barbulescu et al.
(1975) from the Lower Aptian of Baciuliu Rasova (Rumania).

Terebrirostra lyra

 (Sowerby 1816)

Remarks:

 In common with the previous species, the shell is

elongate, but the pedicle tube is comparatively longer in ju-

156                                                                                             GASPARD

veniles. The species is characteristic of the Chloritic Chalk
(Lower Cenomanian) of Cap de la Hève near le Havre
(d’Orbigny coll., no. 6511 and E.M. coll.; Gaspard 1997) and
is also found near Horningsham (England).

Family Kingenidae Elliott 1948, nom. transl.

 Subfamily Kingeninae Elliott 1948, emended Owen 1970

Genus Kingena Davidson 1952

From this subfamily, 4 species are typical for the period:

K. spinulosa

, K. arenosa, K. concinna and K. elegans.

Kingena spinulosa

 (Davidson & Morris 1847)

Remarks:

 This species has been revised by Owen (1970, pl. 2).

The topotypes come from the Upper Albian (Gault)  of Norfolk
and Cambridge. Two specimens were recorded from the Upper
Albian of Orosei (Sardinia) by Dieni et al. (1973). Popiel-Bar-
czyk (1972) recorded specimens from the Middle Cenomanian
near Cracow and also the Lower Cenomanian (?) near Annopol
on the Vistula. The author figured only one specimen with
somewhat crushed hinge-line and anterior  region, but this spec-
imen can be compared to pl. 2, Fig. 6a and pl. 3, Fig. 8 of Owen
(1970) with two sizes of pustules; although crushing prevent to
appreciate the proportions of the shell.

Kingena arenosa

 (d’Archiac 1846)

Remarks:

 This species has been revised by Owen in 1970.

The shell is biconvex, broad, pentagonal, the widest known
among species of the genus with a homogeneous fine pustu-
lation of uniform size, unlike K. spinulosa. The shell exhibits
a great variability, it is generally broader in the posterior
third, with a foramen marked by strong beak ridges. The spe-
cies has been recorded from the Tourtia of Tournai (Tournai,
Montignies-sur-roc, Guissignies (I.R.S.N. coll., no. IG 5694/
1,2), Cherq, Belgium) known to be of Lower Cenomanian
age. It also occurs with Cyclothyris sp., Ovatathyris ovata
(Sowerby), Dereta pectita  (Sowerby) and (?) O. mantelliana
(Sowerby) in Wiltshire (Owen 1970, p. 56).

The species is noted in sandy facies of the Middle Cenoma-

nian in Devonshire and along the Dorset coast as well as in
Middle Cenomanian at Culver Cliff and Compton Bay (Isle
of Wight). It is localized in the Lower Cenomanian and Mid-
dle Cenomanian limestones of Normandy; in the “Essen
Tourtia” Lower Cenomanian of North  Germany; in the envi-
rons of Nagorzanka on the Strypa River (Podolian fauna,
Popiel-Barczyk 1972) and possibly Lower Cenomanian (?)
near Annopol on the Vistula.

Kingena concinna

 Owen 1970

Remarks: 

The shell is broad to oval with a curved umbo, an

anterior commissure generally rectimarginate, and a hinge
line shorter than in K. arenosa. The species was collected
from the Middle Cenomanian (A. rhotomagense Zone), in

the type locality between Folkestone and Dover (Kent). The
paratypes come from Cambridge, Dover, Wiltshire, Lincoln-
shire, Yorkshire. The species is also found in the Boulonnais
(France) in bed 17 of Blanc Nez cliffs with the network of
punctae well exposed.

Kingena elegans

 Owen 1970

Pl. I: Fig. 7

Remarks:

 Species characterized by a pentagonal-elongate

shell, with a hinge-line broader than in K. concinna, as well
as a larger foramen. The shell is marked by regular spaced
growth lines, and a light sulcation affects the brachial valve.
This species characterizes the Turonian (Holaster planus
Zone) (Pl. I: Fig. 7), some specimens coming from the T. lata
Zone (Norfolk).

Numerous specimens, collected by J. Phillip (Univ.

Marseille), between la Bedoule and Cassis (Bouches du Rhône,
France), on the slope (between platform and basin) have consid-
erable affinities with the species, while being smaller than those
from England.

Superfamily Thecidoidea

Remarks:

 Although the superfamily will not be discussed

here, we can refer to the monograph of Pajaud (1970) demon-
strating (p. 270, Fig. 117), the presence of Thecidiopsis Muni-
er-Chalmas (1887), Backhausina Pajaud (1966), Praelacazella
Smirnova (1969) and Vermiculoidea  Elliott (1953) from the
Albian to the Cenomanian.

Conclusion–Discussion

 The stratigraphic distribution of the brachiopod species

presented above is summarised in Table 1. While geographic
distribution and ecological provinces emphasize the fact that
some species are endemic and preferentially confined to spe-
cial environments or facies (such as Sellithyris tornacensis
(d’Archiac) Tourtia of Tournai and Montignies-sur-roc (Bel-
gium), Essen Tourtia (Germany); S. coxwellensis Middle-
miss, Sponge Gravel, (Faringdon, England)), others are cos-
mopolitan or ubiquitous.

(a) Facies and factors of variation

All brachiopod specialists have observed that some species

always tend to be associated with particular sedimentary or
biological facies. This is the case for Cyclothyris, as dis-
cussed by Ager (1965). Also Cyclothyris species with pro-
nounced costae, are found in coarse sediments, or in con-
glomerates (C. scaldisensis  (d’Archiac)). Other brachiopods
associated with coarse sediments appear to be the costellate
terebratellids such as Gemmarcula menardi (Lamarck) and
Arenaciarcula

, whereas the rhynchonellid Grasirhynchia is

found in finer arenaceous sediments. Fine costellate terebratel-

Table 1.

 Occurrence of brachiopod species in NorthWestern Europe between the Aptian and the Turonian (this distribution does not reflect

the totality of the faunas living during these periods). Ammonite biochronozones after J. Thierry et al. (1997). (*= brachiopod markers).

→

DISTRIBUTION AND RECOGNITION IN APTIAN-TURONIAN BRACHIOPOD DEVELOPMENT                         157

158                                                                                             GASPARD

lids such as Dereta pectita (Sowerby) and rhynchonellids:
Grasirhynchia

 and Monticlarella are more frequent in green-

sand, and chalk marls, while representants of the genus Tere-
bratulina

 are often observed in marls.

Moutonithyris moutoniana

 (d’Orbigny), typical of the ear-

ly Tethyan realm, occurs preferentially in clay sediments and
later achieved its invasion northward (Germany, England
(?)). The species is again found in clay and marly lithofacies.
Moutonithyris

 dutempleana (d’Orbigny), evidently a descen-

dant of M. moutoniana (d’Orbigny) is sometimes in the same
context, sometimes in greensand or chalky sediments (Red
Chalk). This fact excludes to find representatives of both Cy-
clothyris

  and Moutonithyris in the same facies.

Specific facies can occur in time, and thus Sellithyris sella

(Sowerby), occurring in marly sediment during the Hauterivi-
an, is more often found in coarser sediments during the Early
Aptian, this fact is in correlation to transgressive movements
and migration routes. On the other hand S. cenomanensis Gas-
pard, a highly biplicate species (with variation of the anterior
commissure) occurring during the Middle Cenomanian, is
found in several lithofacies: in coarse sediments with Cy-
clothyris

 (Sarthe, Indre), in marly sediments (Charentes) or in

chalky-marls (Var). Phaseolina phaseolina (Lamarck) from
the Upper Cenomanian is found in different sediments in rela-
tion to the progression of the marine transgression westward
(see discussion in Gaspard 1985, 1988). Gibbithyris, contrary
to the previous species, occurs mainly in chalky facies.

The distribution presented above shows a great number of

species occurring in the Aptian, albeit not considered in all
the realms (the Jura fauna will be discussed in another pa-
per), and the Cenomanian. These two periods correspond to
large transgressive phases. Particularly in the Cenomanian,
the brachiopods are considered to occur in littoral waters
(Juignet 1974), probably more suitable for Cyclothyris with
sharp costae, with particular environments: Tourtia of Tour-
nai, Essen Tourtia for example. Brachiopod fauna living on
softer sediments (e.g. marls) include genera like Moutonithyris
(probably in deeper conditions), some Sellithyris, generally
smooth-shelled terebratulids and Terebratulina.

The distribution of the different species and their migration

routes already discussed (Middlemiss 1979; Gaspard 1988
mostly for Terebratulids) were dependent upon, and con-
trolled by paleoenvironment: shallow-water and temperature,
high-energy conditions (could be a barrier), argillaceous or
muddy context (poor conditions). In this last context few
genera survive, e.g. Moutonithyris. But morphological adap-
tations to this environment exist: e.g. Terebrirostra species,
(Gaspard 1997). Many species disappear at the end of the
Aptian and Cenomanian periods. Those which appear in the
Upper Aptian seem to be confined mainly to unusual facies
or are endemic (for example species from Faringdon Sponge
Gravel). Numerous species appeared in the Lower Cenoma-
nian such as new species of Sellithyris, Rectithyris depressa,
Capillithyris

  (disparilis and squamosa) and species of Orbi-

rhynchia

, Kingena, Gemmarcula.

(b) Some stratigraphic markers

A preliminary approach in this work, after the presentation

of the paleontological evaluation and distribution, indicates

that some species could be proposed as stratigraphic markers
located according to the standard Ammonite zonation, or
sometimes, the foraminiferal zone. They are denoted by (*)
in Table 1.  These are: Cyclothyris depressa (P. nutfieldensis
Zone),  Gemmarcula crassicosta (Simplorbitolina manasi
Zone, Upper Aptian), Praelongythyris rogeri (L. tardefurcata
Zone), Moutonithyris dutempleana (Stoliczkaia dispar Zone,
M. perinflatum Subzone and basal Hipoturrilites carcitanensis
Subzone), Capillithyris capillata (H. carcitanensis Subzone),
Praelongithyris fecampi

 (H. carcitanensis & Mantelliceras

saxbii Subzones), Monticlarella brevirostris (upper-half of
Mantelliceras cantianum and Lower part of M. dixoni Sub-
zones), Cyclothyris difformis (part of M. dixoni Subzone and
Middle Cenomanian), C. lamarckiana (Marnes à O. biauricu-
lata

,  Calycoceras  naviculare),  C. compressa  (C. naviculare,

base of Upper Cenomanian), Grasirhynchia martini (upper
part of Lower Cenomanian), Modestella geinitzi and Kingena
concinna

 (lower part of Acanthoceras rhotomagense Zone),

Orbirhynchia mantelliana

 from M. dixoni Zone to Turrilites

acutus–T. costatus Subzones boundary in A. rhotomagense
Zone), Concinnithyris subundata (T. acutus Subzone and part
of A. jukesbrownei Zone), C. obesa (Middle Cenomanian),
Gemmarcula menardi 

(A. juskesbrownei Zone), Phaseolina

phaseolina 

(part of C. guerangei to upper part of Neocardio-

ceras judii Zones), Orbirhynchia boussensis-wiesti-cuvieri
group (N. judii Zone to basal I. labiatus Zone), Concinnithyris
protobesa

 (Lower Turonian), Terebratulina lata (Collignon-

iceras woolgari Zone=T. lata Zone of the English workers),
Kingena elegans 

(H. planus Zone and (?) T. lata Zone). Creti-

rhynchia

 and Gibbithyris would certainly also supply some

complementary Turonian markers.

(c)  Presence of these faunas elswhere in the world during
the Aptian-Turonian periods

Before the discussion concerning a proposal for the recognition

of phases in the brachiopod development during the Aptian–Turo-
nian periods in NW Europe, it would be interesting to have a look
at the presence, elsewhere, of some genera and species previously
listed. The comparison is not easy because the Cretaceous is not
well exposed worldwide, and even where it is exposed brachio-
pod specialists are not always on land to describe the faunas.

Some indications come from North America, Madagascar,

China, Caucasus and the Crimea.

Representatives of Moutonithyris, a mainly subtethyan genus,

are manifest also in North Africa, Madagascar (Collignon 1949,
1950), and the Caucasus, the Crimea and North America, partic-
ularly Central Mexico (Chiodi et al. 1988). Consideration of the
brachiopod present in Canada and Mexico and North America is
generally given by Sandy (1990a,b).

Sellithyris

 and Praelongithyris are also present in the Cau-

casus and the Crimea (Smirnova 1972) and in the Crimean
region, Klikushkin (1973) referred to species of Lingula
from Upper Cenomanian.

Sellithyris

–Praelongithyris assemblages are known also in

the limestones and marls of the Lower part of Langshan For-
mation, Northern Xizang (Sun 1987), but these are composed
mainly of new species which need close comparison with
European species. For example, Sun proposed a parallel be-
tween  Praelongithyris rogeri Gaspard and P. xiongmeinsis

DISTRIBUTION AND RECOGNITION IN APTIAN-TURONIAN BRACHIOPOD DEVELOPMENT                         159

Sun or between Sellithyris sella (Sowerby) and S. pentago-
naris 

Sun. Also an Orbirhynchia-Parakingena assemblage

occurs in the limestones exposed near North Baingoin Lake,
which are expected to be of Late Albian age by Sun.

Some Capillithyridinae are present in the South of URSS

(Lobacheva 1986), most of the genera quoted in the study are
also listed in Smirnova (1990). Some species are equivalent
to those in Europe: Moutonithyris moutoniana, Loriolithyris
russillensis

,  Sellithyris sella, Praelongithyris praelongifor-

ma

, Rhombothyris extensa, Cyrtothyris cyrta, Rectithyris de-

pressa

. If all these genera are represented, their species con-

tent differences and some representatives will need
investigative work to be critically sure of it synonymy. Col-
laboration is needed to improve the  genera and species
stratigraphical distribution.

 (d) Recognition of phases in the brachiopod development
       — A proposal

In addition to the appreciation of some markers, the above

distribution summarised in Table 1 allows to approach the
recognition of several phases in the brachiopod development
for the period considered. These phases correlate with trans-
gressions and regressions, their reality in time depends main-
ly upon the state of published data and is susceptible to be
progressively improved with new supplies in stratigraphical
distributions.

The proposal can be presented as follows:
“Phase A” could  cover the Lower Aptian with many rep-

resentative genera. The impact of the Northern transgression
results in the differentiation of species approximately at the
boundary of Cheloniceras martinioides and Parahoplites nut-
fieldensis Zones. The main genera recorded are: Sellithyris,
Musculina

, Moutonithyris, Loriolithyris.

“Phase B” beginnning approximately  with the P. nutfielden-

sis Zone includes representatives of Cyclothyris, Praelon-
githyris

, Platyhyris, Gemmarcula  (crassiscosta and aurea),

Rhombothyris

,  Sulcirhynchia,  Burrirhynchia and Lingula

truncata

.

“Phase C” corresponds to the extinction of species origi-

nating from the Aptian trangression. Some differentiations in
fauna were apparently due to north-westward migration in
the Paris Basin, or contrary to the Tethys.

“Phase D” beginning approximately at the limit of Leymeri-

ella tardefurcata and Douvilleiceras mamillatum Zones is a
quiet period without a great explosion of species during all the
Middle Albian and mainly the Lower part of Upper Albian.

“Phase E” beginning approximately at the Albian-Cenom-

anian boundary  presents a rich fauna in relation to the trans-
gressive expansion, the top of which is highlighted by the up-
per limit of Orbirhynchia mantelliana.

“Phase F”: during the Cenomanian, a second proliferation

in the development of some species is characterized at the
T. costatus-T. acutus Subzones boundary, by the differentia-
tion of some Concinnithyris and Gibbithyris  species which
continue to evolve, by Grasirhynchia, Cyclothyris and
Modestella

 for instance. This phase ends with Phaseolina,

Orbirhynchia

 boussensis-wiesti, Gemmarcula carentonensis.

“Phase G” at the Cenomanian-Turonian boundary (O. cu-

vieri

) concerns the expansion of Concinnithyris, Gibbithyris,

Cretirhynchia

 in more chalky facies.

As a proposal these phases are not static and more precise data

during material sampled worldwide in the future will be usefull.

Acknowledgements:

 The author would like to express many

thanks to the Crurators of the different collections mentioned
and to the anonymous reviewers.
** This is a contribution to UMR 5561 and to IGCP Project
no. 362 (Tethyan and Boreal Cretaceous).

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