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THE NATU
| HISTORY MUSEUN

12
} PRESENTED

Geology Series

HISTORY
MUSEUM

VOLUME 50 NUMBER 1 23 JUNE 1994

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

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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Geol.)

Geology Series
ISSN 0968-0462 Vol. 50, No. 1, pp. 1-104
The Natural History Museum
Cromwell Road
London SW7 5BD Issued 23 June 1994

Typeset by Ann Buchan (Typesetters), Middlesex
Printed in Great Britain at The Alden Press, Oxford

Bull. nat. Hist. Mus. Lond. (Geol.) 50(1): 1-103 Issued 23 June 1994

Systematics of the melicerititid cyclostome
bryozoans; introduction and the genera Elea,
Semielea and Reptomultelea

PAUL D. TAYLOR
Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK

CONTENTS
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P. D. TAYLOR

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AAAAA

Synopsis. The Family Eleidae, whose species are commonly referred to as melicerititids, is an unusual clade of
cyclostome bryozoans with operculate zooids, a homoplasy shared with cheilostomes. Melicerititids range from the
Barremian to the Palaeocene and are mostly European in geographical distribution. They are especially abundant
and diverse in some Cenomanian-Campanian deposits of England, France and Germany. This paper is the first part
of an intended comprehensive systematic revision of melicerititid species. Following a brief account of the history of
melicerititid research and a review of their morphology, the main part of the paper describes 36 species belonging to
the three fixed-walled genera (Elea, Semielea and Reptomultelea) with lamellar colony forms. Almost all of these
species have been studied with the aid of scanning electron microscopy, and most of the type material has been
re-examined and lectotypes chosen when necessary. The following 20 species are new: Elea elegantula, E. flabellata,
E. mackinneyi, E. pseudolamellosa, E. subhexagona, E. viskovae, E. whiteleyi, Reptomutltelea acclivata, R. auris, R.
bituberosa, R. convexa, R. goldfussi, R. levinseni, R. matutina, R. mitrus, R. parvula, R. pegma, R. pseudopalpeb-
rosa, R. reedi and R. scanica. Reptomultelea betusora is proposed as a new name for R. tuberosa (Reuss), a junior
homonym of R. tuberosa d’Orbigny. Keys are provided for the identification of melicerititid genera, and for species

of Elea and Reptomultelea.

INTRODUCTION

This is the first of a planned series of papers intended to
provide a complete systematic account of the cyclostome
bryozoan family Eleidae, commonly referred to as ‘melicer-
ititids’ (Family Melicerititidae) and occasionally as ‘opercu-
late cyclostomes’. Melicerititids range from the Barremian
Stage of the Lower Cretaceous to the Danian Stage of the
Palaeocene, a duration of over 60 MA. Almost all species
come from localities in Europe; melicerititids are presently
unrecorded from the Americas, Africa, Antarctica and Aus-
tralasia. They are important constituents of many bryozoan
faunas, especially in the Upper Cretaceous of western
Europe. However, melicerititids have never been compre-
hensively monographed, although Levinsen (1912) did revise
many of the commoner species in the only major publication
devoted to the group.

Melicerititids were chosen for detailed study for three
principal reasons:

1. They clearly constitute a monophyletic group (clade),
defined by the possession of a calcified zooidal operculum,
and are therefore one of very few unequivocal clades which
can be currently distinguished among cyclostome bryozoans
or indeed stenolaemates in general. Questions concerning
evolutionary patterns can be addressed with more clarity in
monophyletic groups (see Smith, in press).

2. Melicerititids have a greater number of morphological
characters for use in taxonomy than most other cyclostomes,
principally because of the wide variety of apertural shapes
and zooidal polymorphs they possess.

3. The operculum and avicularium-like polymorphs of
melicerititids evolved in parallel with those of contemporane-
ous cheilostomes, providing an opportunity for comparative
study of morphological and taxonomic diversification in two
distantly-related groups of bryozoans.

These factors mean that melicerititids are a good target
group among stenolaemate bryozoans for studies of evolu-

tionary patterns; for example, evolutionary trends, conver-
gent evolution, and patterns of extinction and radiation.

HISTORY OF RESEARCH

The early history of research on melicerititids was fully
chronicled by Levinsen (1912) and will only be summarized
here. The first melicerititid species to be formally named was
Ceriopora gracilis Goldfuss, 1826, from the Cenomanian of
Essen in Germany. This species was subsequently assigned by
Roemer (1840) to Meliceritites, the first genus to be proposed
for a melicerititid. Roemer also assigned to his genus two
other Cretaceous species, Ceriopora roemeri v. Hagenow,
1839 and a new species, Meliceritites porosa. The genus name
Meliceritites was derived from the cheilostome genus Melic-
erita on account of their superficial similarity, notably in the
hexagonal frontal shapes of the zooids.

The history of higher classification of the melicerititids
began in 1851 when v. Hagenow referred Meliceritites (as
Escharites) and Inversaria (now known to be a cheilostome,
see Voigt & Williams, 1973; Voigt, 1974) to his division
Salpingina, claiming to have identified opercula in both
genera. Levinsen (1912) was probably correct in doubting v.
Hagenow’s claim: one of the species (Escharites [Filicea]
velata v. Hagenow) in which opercula were supposedly seen
by v. Hagenow lacks opercula but has terminal diaphragms
that could have been mistaken for opercula. A year later
d’Orbigny (1852) founded a division — Centrifuginés opercu-
linés — for two of his new families, Eleidae and Myriozoumi-
dae, making clear from the diagnosis that he had observed
true opercula. While the Eleidae is nowadays the accepted
family for Meliceritites and related genera of operculate
cyclostomes, including the type genus Elea, the Myriozoumi-
dae is recognized as a family of ascophoran cheilostomes.

Hamm (1881) included Meliceritites, together with two
non-operculate cyclostomes (Stigmatopora Hamm and Cyrto-

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 3

pora Hamm), in the Stigmatoporina, a highly artificial divi-
sion not adopted by later workers. Another disregarded
group is the “Typus’ Metopoporina under which Marsson
(1887) united the Eleidea [sic] and Ceidea [sic] based on their
trumpet-shaped zooids (the remaining cyclostomes were
placed in the ‘Typus’ Solenoporina). Despite the disuse of
Metopoporina, the practice of grouping together these two
families has persisted, notably in the Treatise (Bassler, 1953).
Pergens (1890) referred the Eleidae to his Melicertitina.

Whereas Hamm (1881), Marsson (1888) and Pergens (1890)
denied the presence of an operculum in melicerititids, Waters
(1891) thought that only a chitinous operculum might have been
present. Waters did, however, recognize the similarity between
the eleozooids of melicerititids and cheilostome avicularia.
Gregory (1899) confused opercula with terminal diaphragms
(like several of his predecessors), but acknowledged the exist-
ence of ‘avicularia’, and observed brood chambers (‘gonoecia’
and ‘gonocysts’) in melicerititids. He also considered ceid cyclos-
tomes to be merely worn specimens of melicerititids in which the
frontal wall has been lost.

D’Orbigny’s (1852) recognition of opercula was reaffirmed by
Levinsen (1902). Lang (1906) very briefly revised the encrusting
melicerititids, but his key to their generic identification is flawed
in several respects. Thirty species and three varieties of melicer-
ititids were described by Levinsen (1912) in a short monograph
remarkable for the accuracy of its detailed observations and
careful interpretations. Few advances have been made in under-
standing melicerititids since Levinsen’s fundamental paper,
although some papers have included systematic descriptions or
figures of melicerititid species, notably those of Canu & Bassler
(1922, 1926), Voigt (1924, 1928, 1951, 1953, 1960, 1962, 1967,
1973, 1975a, 1975b, 1981, 1983, 1985a, 1985b, 1989), Prantl
(1938), Viskova (1965, 1970), Walter and coauthors (Walter
1975, 1977; Delamette & Walter, 1984; Masse & Walter, 1974;
Walter & Clavel, 1979; Walter et al., 1975), Brood (1972), Taylor
(1987a, b), Pitt & Taylor (1990), and Favorskaja (1992).

Aspects of melicerititid morphology and palaeobiology
were treated by: Boardman (in Boardman et al., 1983), who
reconstructed the soft tissues around the operculum; McKin-
ney (1975), who described zooecial budding patterns of
melicerititids and other dendroid stenolaemates; Taylor
(1982), who described probable predatory borings in melicer-
ititid zooids; and Taylor (1986a), who reviewed polymor-
phism in the group. More recently, Schafer (1991) has
described the gonozooids of melicerititids, and Taylor (1990)
used melicerititids as an example of the application of scan-
ning electron microscopy in bryozoology.

The systematic position of melicerititids within the Cyclos-
tomata has received scant attention. Bassler (1953), in the
bryozoan Treatise, grouped the Eleidae with the Ceidae (=
Semiceidae Buge, 1952), placing the two families in the
Suborder Salpingina. Viskova and Morozova (1988) and
Viskova (1992) published an unorthodox classification which
recognized three post-Palaeozoic orders of stenolaemates:
Tubuliporida Blainville, Cerioporida Bronn, and Melicer-
ititida Pergens. Within the Order Melicerititida they placed
the families Eleidae d’Orbigny, Melicerititidae Pergens,
Semiceidae Buge and Lobosoeciidae Canu & Bassler. There
is little evidence to suggest a closer relationship between
melicerititids and the two latter families than between melic-
erititids and several other tubuliporine families. Further-
more, partitioning the melicerititids between two families
(Eleidae and Melicerititidae) rather than uniting them in a
single family (Eleidae) implies an understanding of relation-

ships between genera which is currently lacking; for example,
it is unclear whether Reptomultelea (even if monophyletic) is
more closely related to Elea than to Meliceritites (see p. 47).

General morphological comparison suggests that the sister
group of melicerititids probably lies among non-operculate
tubuliporine cyclostomes, notably Collapora and its relatives,
which are traditionally assigned to the Family Multisparsidae
(= Macroeciidae). Pending a comprehensive phylogenetic
study, the Eleidae are here placed within the Suborder
Tubuliporina Busk, 1852, although it is acknowledged that
this suborder as currently understood is almost certainly
paraphyletic.

MORPHOLOGY

The principal morphological characters observed in melicer-
ititids are summarized in Table 1. Key features of melicer-
ititid morphology are discussed below, with important
terminology printed in bold, and the external appearance of
zooidal polymorphs is depicted diagrammatically in Figure 1.
It must be emphasized that the text below is not intended as a
comprehensive account of all of the morphological variations
found in melicerititids.

Colony-form

Despite the unusual morphology of the zooids in meliceritit-
ids, the range of colony-forms within the family is typical of

small eleozooid
(demizooid)

intramural
demizooid

autozooid with
terminal diaphragm

autozooid with
operculum

autozooid with
open aperture

large eleozooid
(rostrozooid)

kenozooid

intramural
autozooid

Fig. 1 Diagrammatic illustration of external zooidal morphology
and polymorphism in melicerititid cyclostomes.

Table 1 Principal morphological characters of melicerititids.

Colony

a. colony-form (encrusting, multilamellar, vinculariform without
axial canal, vinculariiform with axial canal, adeoniform,
eschariform, cavariiform)

b. branch width/colony layer thickness

c. overgrowths (presence, abundance, polymorph type of
pseudoancestrula)

d. base (extent, ancestrula, zone of astogenetic change)

e organization (fixed-walled, free-walled, mixed)

f. zooid arrangement (whorls, quincunx, irregular, one side of

branch only)

Autozooids

frontal length

frontal width

frontal elongation (length:width ratio)

frontal shape

boundary wall

aperture length

aperture width

aperture elongation (length:width ratio)
aperture shape

size of aperture relative to frontal area
apertural shelf

hinge teeth/bar

abundance of in-situ opercula

opercular pseudopore number and distribution
surface ornament of opercula

opercular sclerites

terminal/subterminal diaphragms (abundance, placement,
pseudopores, perforations)

18: intramural buds

OVORR OAT OER moe aN se

Eleozooids

a._r. of autozooids, and:

S. affect on surrounding az (e.g. overgrowth, displacement)
lic rostral platform

Kenozooids and cancelli
a. distribution
b. surface features

Gonozooids

abundance

total frontal length
dilated frontal wall length
frontal width

frontal shape

inflation

ooeciopore length
ooeciopore width
ooeciopore shape (length: width ratio)
atrial ring

floor morphology

Foro moe ae oe

that encountered in many cyclostome groups. The following
main colony-forms occur, in approximate order of frequency:
erect dendroid (Figs 3, 5—7), encrusting multiserial and
multilamellar (Figs 2,4, 181-184), erect bifoliate (Figs 17, 49,

‘P. D. TAYLOR

87), erect tubular (Figs 110, 119, 121-123) and erect fenes-
trate (see Taylor 1987a). Common growth-forms of cyclos-
tome bryozoans not represented among melicerititids are
uniserial or pauciserial (ribbon-like) encrusting colonies, and
articulated erect colonies.

All colonies begin with an encrusting base, usually attached to
a hard substrate such as a shell (Figs 37, 134, 211) or pebble.
However, some colonies were evidently attached to a perished
organic substrate, leaving a mould bioimmuration usually in the
shape of a hollow cylinder. In some cases, the edge of the original
substratum was overlapped and the younger parts of the colony
base grew freely over the sea-bed, becoming essentially free-lying
(Fig. 183). Secondary substrates were occasionally incorporated
into the bases of these free-lying colonies as they expanded across
the sea-bed.

Erect growth is entirely lacking in species of Reptomultelea
and the exclusively encrusting colonies are typically multila-
mellar (Fig. 4). However, the majority of melicerititid species
developed erect growth from an encrusting base. Most erect
colonies have subcylindrical branches, less than 2 mm in
diameter, which bifurcate to give a dendroid, bushy colony
(Figs 5-7) equivalent to the vinculariiform morphotype of
cheilostomes. Species of Meliceritites, Atagma, Meliceritella
and Foricula all have this colony-form. In most of these
colonies, feeding zooids are evenly distributed around the
circumference of the branches which, therefore, fall within
the radial non-maculate category of McKinney (1986a, 5).
However, a few species with wider branches may develop
raised monticular maculae (e.g. Foricula aspera). Multiple
lamellar overgrowths are characteristic of many species with
erect branches, particularly fixed-walled dendroid and broad
bifoliate colonies. Branches of these colonies resemble the
Jurassic cyclostome Terebellaria (Taylor, 1978) in cross-
section, and may be termed ‘terebellariiform’. In Meliceritella
autozooids have a restricted distribution around the circum-
ference of the subcylindrical branches which have a well-
defined reverse or dorsal side composed of kenozooids and/or
eleozooids. This colony morphotype therefore falls within the
unilaminate category of McKinney (1986a, b). Meliceritella
specimens are invariably recovered as short broken branches
and it is usually uncertain whether colonies were originally
bushy and three-dimensional or were almost two-dimensional
planar fronds. However, scarcity of branch anastomoses
suggests that most colonies were bushy with the exception of
one species, M. schneemilchae, in which regular branch
bifurcation and anastomosis in a single plane gives a fenes-
trate colony-form (Taylor, 1987a).

Less common among melicerititids are dendroid erect
colonies with bilaminate branches in which zooids bud from
both sides of a median lamina (Figs 26-27). Bilaminate
colonies can have either narrow, strap-like branches which
bifurcate in the plane of the budding lamina (e.g. Elea
elegantula), or broad, foliaceous fronds which can be folded
into complex corrugations or anastomosed into box-like
structures (e.g. E. lamellosa). These occur in the fixed-walled

Figs 2-7 Colony forms in melicerititid cyclostomes. 2, multilamellar encrusting colony of Reptomultelea sarthacensis (d’Orbigny, 1853); EM
RE 551.763.31.A711, Cenomanian, Essen, Germany; x 1-4. 3, typical assemblage of fragments of erect dendroid species; BMNH sample,
Santonian, Craie de Villedieu, Villedieu, Loir-et-Cher, France; ca x 3. 4, worn edge of multilamellar colony of Reptomultelea sp. showing
layering; EM RE 554. 763. 31. A745/9, Cenomanian, Essen, Germany; X 4-5. 5, erect dendroid colony of Meliceritites dendroidea
(Keeping, 1883) embedded in a matrix of glauconitic sandstone; BMNH D3145, Aptian, Lower Greensand, Shanklin, Isle of Wight,
England; x 0-8. 6, dendroid specimen of M. dendroidea with a densely-branched colony-form; BMNH D55269, Aptian, Faringdon Sponge
Gravel, Little Coxwell Pit, Faringdon, Oxfordshire, England; x 2-2. 7, small, loosely-branched dendroid colony of M. semiclausa
(Michelin, 1846) with a concave encrusting base of small area; BMNH D3695, Cenomanian, Le Mans, Sarthe, France; x 4-4.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

FaSeUiT ies BNP es
iactsaceecege
Ba

BEGAN back

genus Elea and its free-walled analogue Biforicula.

Consistently tubular branches characterize the genus
Semielea. The axial lumen (Fig. 119) of these branches is
about 1 mm in diameter, and lined by an apparent exterior
wall which is transversely wrinkled and whose origin can be
traced back to the basal lamina of the colony base (Fig. 122).
These ‘cavariiform’ branches seem not to be the result of
growth around a perished substratum as the walls of the
lumen evidently grew freely into space and do not bioimmure
a substratum. Furthermore, the lumen is occasionally divided
by transverse platforms which could not have formed in a
colony wrapped around a cylindrical substratum (cf. Lophol-
epis radians, see Voigt, 1982: pl. 9, figs 3-6).

Much narrower axial canals occur in some species of
Meliceritites (e.g. M. semiclausa; see McKinney, 1975: pl. 3,
fig. 3a) but appear to be formed of interior rather than
exterior wall. These resemble the axial canals found in the
Mesozoic non-melicerititid tubuliporine genera Entalophora
(see Walter, 1970), Bicoronipora (see Walter, 1987), and
Coelospiropora (see Voigt, 1968; Voigt & Flor, 1970; Walter,
1987), and in the late Palaeozoic cryptostome genus Rhab-
domeson (see Blake in Boardman et al., 1983). Axial canals
of this type are of uncertain origin and function. Some may be
greatly-elongated polymorphic zooids. It is possible that axial
canals provided a pseudocoelomic connection of physiologi-
cal value between branch growing tips and the older, more
proximal zooids of the colony. Alternatively, their function
may have related to the provision of a surface from which
zooidal budding could occur.

Overgrowths can be found in most melicerititid species.
They usually originate by intrazooecial fission (sensu Hillmer
et al., 1975) and subsequent eruptive budding of the newly-
formed zooids onto the colony surface. Incipient overgrowths
are first evident as apertures divided into an average of six
chambers by radial walls arranged as spokes around an inner
chamber enclosed by a ring-like wall (Figs 250, 271-272, 284).
The inner chamber is apparently continuous with that of the
parent zooid and develops into a pseudoancestrula at the
centre of the overgrowth (Figs 131, 163, 179, 206-207,
212-213, 221, 225, 285). The pseudoancestrula is encircled by
radially-orientated zooids originating from the surrounding
chambers. A circular overgrowth with a circumferential grow-
ing edge is thus formed. Often several closely-spaced over-
growths are present and these coalesce as they grow outwards
and come into contact with one another. Overgrowths are
structurally distinct units and can be classified as subcolonies.
Conspicuous secondary zones of astogenetic change are
developed, beginning with the pseudoancestrula, which has a
small aperture and little or no frontal wall. Aperture and
frontal wall dimensions increase progressively through the
succeeding generations of zooids. In species with high and
pointed apertures (e.g. Meliceritites gracilis (Goldfuss)), zoo-
ids in secondary zones of change also tend to have propor-
tionally shorter and more rounded apertures than zooids
from zones of astogenetic repetition.

Skeletal organization

The major structural walls of bryozoan colonies can be
categorized as basal walls, vertical walls and frontal walls. In
all cyclostome bryozoans basal walls are apparently exterior
walls, secreted from one side only, and include a calcified
layer between the secretory epithelium and cuticle. Vertical
walls and frontal walls may be with or without calcified

P. D. TAYLOR

layers. Three basic skeletal organizations (see Taylor &
Larwood, 1990: fig. 10.6 for a simplified representation, and
Boardman in Boardman et al., 1983 for more detailed expla-
nations of the latter two organizations) can be recognized
according to which of these walls have calcification:

1. Corynotrypid. Known only in the Palaeozoic Family
Corynotrypidae, this organization is characterized by calci-
fied frontal exterior walls but non-calcification (or absence)
of vertical walls.

2. Fixed-walled. Previously termed single-walled (Borg,
1926), fixed-walled organization has calcification of both
interior vertical walls and exterior frontal walls. Frontal walls
are fixed to the ends of the vertical walls.

3. Free-walled. Previously termed double-walled (Borg,
1926), here only the interior vertical walls are calcified. The
frontal exterior wall remains uncalcified and is free of the
vertical walls, enclosing a hypostegal pseudocoel between
itself and the ends of the vertical walls.

Skeletal organization has been employed in the subordinal
division of cyclostomes: two suborders (Tubuliporina and
Articulata) are essentially fixed-walled, and three suborders
(Cerioporina, Cancellata and Rectangulata) essentially free-
walled. However, it is becoming increasingly clear that the
taxonomic distinction between fixed- and free-walled cyclos-
tomes is more complex. For example, the gonozooids of
otherwise free-walled cerioporines are fixed-walled (e.g.
Schafer, 1991); individual branches of Cinctipora elegans can
show alternations of free- and fixed-walled autozooids
(Boardman et al. , 1992); and Boardman (1975) has described
fixed-walled organization in a species of the normally free-
walled genus Heteropora.

Melicerititids provide another example of the mixing of
skeletal organizations. In all species for which they have been
described, gonozooids are fixed-walled, whereas autozooids
(and eleozooids) can be either fixed- or free-walled. Two
melicerititid genera (Foricula and Biforicula) have free-
walled autozooids, but the remaining genera possess fixed-
walled autozooids. The areas between autozooidal apertures
in free-walled genera are occupied by cancelli (Fig. 8). In
contrast, the calcified frontal walls of the autozooids them-
selves occupy these areas in fixed-walled species (Fig. 8).
Regardless of whether the free- or fixed-walled organization
is the more primitive (see the inconclusive discussion in
Boardman et al., 1992), the existence of both organizations in
the demonstrably monophyletic melicerititids shows that at
least one of the organizational types must be polyphyletic in
cyclostomes as a whole.

Zooid structure

Melicerititids are notable among cyclostome bryozoans for
their high levels of zooidal polymorphism (Taylor, 1986a).
Polymorphism is defined as discontinuous variation in mor-
phology between zooids within a colony, and is known or
inferred to reflect differences in function between zooids (see
Boardman & Cheetham, 1973). Although different polymor-
phs are recognized morphologically, their naming in both
Recent and fossil bryozoans is usually based on presumed
function rather than homology (see Silén, 1977). This can be
a difficult task in fossil bryozoans. A fundamental division is
made between feeding zooids, termed autozooids, and non-
feeding zooids, termed heterozooids. Sometimes two auto-
zooidal polymorphs are present, for example the A- and
B-zooids of the cheilostome Steginoporella (Banta, 1973).

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 7

aperture

frontal wall

fixed-walled organization

aperture

@©O
: e<_ cancelli

“ © ‘
free-walled organization

Fig. 8 Diagrammatic illustration comparing fixed-walled and
free-walled organizations in melicerititid cyclostomes.

Heterozooids in modern cyclostome bryozoans include gono-
zooids (sometimes called gynozooids to avoid terminological
and functional confusion with androzooids; Silén, 1977), in
which larvae are brooded, and kenozooids which is a ‘waste-
basket’ category of atrophied zooids seemingly having space-
filling roles. Both of these heterozooid types can be
distinguished with relative ease in melicerititids as they differ
little from the same polymorphs in modern species of cyclos-
tomes.

Most melicerititids have an additional group of polymorphs
characterized by opercula and apertures which are modified
relative to other zooids in the colony. These have been
termed eleozooids by Taylor (1986a), a derivative of Canu’s
(1897a) term ‘eleocellaires’.

Internal morphology

Like most stenolaemates, the proximal parts of the zooids in
endozones are long, narrow, thin-walled tubes orientated
subparallel to growth direction (Fig. 9). With growth they
gradually diverge from the branch axis and enter the exozone.
At the endozone-exozone junction, zooids rapidly increase in
width, their walls thicken, and they bend to become orien-
tated almost perpendicular to the branch surface. Differences
between melicerititid polymorphs generally only become
clear in the exozone and are much more apparent on the
colony surface than in tangential or other sections cut through
colonies.

Transverse sections of melicerititid branches are notable
for the strong contrast between the abundance of small
diameter zooidal tubes in the endozone and the fewer large

Figs 9-10 Thin sections of a typical dendroid melicerititid,
Meliceritites palpebrosa Levinsen, 1912; USNM 2634-16,
Coniacian [?Santonian], Villedieu, Loir-et-Cher, France; 9,
longitudinal section showing long, club-shaped zooids originating
in the endozone, X 16; 10, transverse section showing small
diameter of zooids in the endozone and spiral arrangement of
buds becoming evident at the endozone-exozone transition, x 40.

zooidal tubes in the exozone (Fig. 10). Budding of zooidal
tubes occurs entirely in the endozone, with the exception of
some very small polymorphs (e.g. demizooids in Afagma) and
the cancelli of Foricula and Biforicula, which may bud in the
shallower parts of the exozone close to the colony surtace.
Sometimes the zooids are arranged in spiralling rows in the
outer endozone and exozone (McKinney, 1975; Fig. 10). At
the bases of erect colonies and in unilamellar and multilamel-

lar colonies, new buds arise by division of vertical interior
walls at their junction with a basal budding lamina of exterior
wall. Bilamellar colonies have a similar style of budding but
here the budding lamina is an interior wall (Figs 26-27).
Dendroid colonies with axial canals may display budding
around the periphery of the canal (McKinney, 1975: pl. 3, fig.
3a).

Intrazooidal structures are few within the endozone. Occa-
sional thin diaphragms have been observed in some species
(e.g. basal diaphragms in Foricula spp.), and mural spines are
present deep within the endozone in zooids of Elea triangu-
laris (Michelin) (Fig. 93). However, it should be noted that
only a small minority of species have been studied in thin
section and it is possible that internal structures are as yet
undetected in other species.

Early astogeny

Early growth stages are known in very few species. While the
bulk of melicerititid specimens are fragments from distal
parts of colonies, those specimens which do preserve proxi-
mal parts generally have them completely obscured by the
overgrowth of later zooids. Nevertheless, a few examples
have been found of ancestrulae and succeeding zooids from
the primary zone of astogenetic change. In all cases the
ancestrula has a large protoecium (= primary disc) and a
comparatively short distal tube (e.g. Fig. 288). This contrasts
with most tubuliporine cyclostomes in which the distal tube is
appreciably longer than the protoecium. Opercula have not
been observed in-situ, but the straight proximal edge of the
aperture strongly suggests that the ancestrula was operculate
like the zooids of later astogenetic stages. The primary zone
of astogenetic change shows the progressive increase in zooid
size which is typical of early colony growth in bryozoans.
Unfortunately, early astogeny is unknown in free-walled
melicerititids and the presumed astogenetic transition
between fixed- and free-walled organization has yet to be
elucidated (cf. Cinctipora elegans as described by Boardman
et al., 1992).

Autozooids

The outline shape of melicerititid autozooids varies between
species but most often approximates a longitudinally elon-
gated hexagon with two longer sides parallel to the long axis
of the zooid (Fig. 11). Less commonly, autozooidal outline is
‘thomboidal’, the proximal and distal sides at 90° to the long
axis (Fig. 11). Four-sided, roughly diamond-shaped autozoo-
ids may also occur locally within colonies, apparently as
derivatives of the hexagonal pattern in which the lateral sides
have been eliminated. The distal angles of hexagonal zooids
are variably rounded so that they are subparallel to the
distolateral edge of the aperture, which is always positioned
close to the distal end of the zooid.

Fixed-walled species have flat to slightly convex frontal
walls pierced by pseudopores (by analogy with modern
fixed-walled species, pseudopores would have been cuticle-
covered during life). Whereas pseudopore shape varies con-
siderably in other cyclostome groups, pseudopores in
melicerititids are always circular; slit-shaped, teardrop-
shaped and gull-shaped are unknown in melicerititids. The
density of pseudopores characteristically declines approach-
ing the apertural hingeline.

Zooidal boundaries vary in prominence in fixed-walled

P. D. TAYLOR

hexagonal zooids

'rhomboidal' zooids

Fig. 11 Comparison between melicerititid cyclostomes with zooids
of hexagonal and ‘rhomboidal’ surface outlines showing
differences in the packing and arrangement of apertures.

species of melicerititids. Sometimes they are defined by a
slightly salient wall lacking pseudopores, but in other
instances pseudoporous frontal wall appears to extend unin-
terrupted across the boundary from one zooid to the next.
Low, rounded tubercles may be formed from raised patches
of zooidal boundary wall at the angles of the frontal walls in
some species. Their location in hexagonal autozooids gener-
ally corresponds approximately to the two ends of the aper-
tural hingeline and the distal tip of the aperture. In Elea
labyrinthica (Michelin), a prominent tubercle is situated
distally of the aperture (Fig. 61).

Autozooid apertures in fixed-walled species are positioned
more-or-less terminally — no or very little frontal wall is
present distally of the aperture. The shape of the aperture
varies between species but is relatively constant within a
species and provides an extremely useful character for species
recognition (Figs 14, 125). Most species have apertures which
are longer than wide, some have equidimensional apertures,
and a few have apertures wider than long. The proximal edge
of the aperture — the hingeline — is straight or slightly convex
distally (bowed). The distal edge can be well-rounded, sub-
rounded, sub-angular, or distinctly pointed. As a result,

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 9

apertural shapes found in melicerititids generally range from
a semicircle at one extreme to a gothic arch at the other.
More complex shapes are occasionally found, for example,
the ogee-arch shaped apertures of Reptomultelea sarissata
(Figs 282-283).

Hinge lines when well-preserved usually have a pair of
small teeth or condyles, often with a ridge extending between
them. Failure to observe these hinge teeth in some species is
more likely to reflect preservational deficiency than true
absence. Meliceritites divergens (d’Orbigny) is notable both
for the large size of its hinge teeth and the presence of
additional ‘drop-like’ teeth around the distal edge of the
aperture (cf. the beading found on the distal edge of aper-
tures in the cheilostome Rhynchozoon). Apertural shelves
(e.g. Fig. 251) extend around the distal margins of the
autozooidal apertures in many species of melicerititids and
apparently form platform on which the closed operculum
rested. Again, these vary in development between different
species, ranging from barely discernable narrow shelves
which are typically developed only in the distalmost parts of
the aperture, to broad shelves of equal width along the entire
distal margin of the aperture.

Compared with other cyclostomes, peristomes in melicer-
ititids are extremely short and cannot readily be distinguished
as a separate skeletal unit of the zooid. This contrasts with the
long peristomes in Plagioecia patina and other tubuliporines
which are marked by their sparse pseudopores and which may
be shed as a unit during late ontogeny (see Silén & Harmelin,
1974). In the great majority of melicerititid species no more
than a slightly raised apertural rim is developed. However,
Meliceritites dollfusi Pergens has opercula located atop rela-
tively well-defined but short peristomes.

Terminal diaphragms can be found in most melicerititid
species but their distribution within colonies is generally
without obvious pattern; seldom is there a clear ontogenetic
zonation with terminal diaphragms in only the older, more
proximal zooids, as found in many other cyclostomes (Silén &
Harmelin, 1974; Boardman et al., 1992). A variety of differ-
ent types of terminal diaphragms can be distinguished, occa-
sionally with more than one type being present in a single
colony (or species). Some terminal diaphragms are
pseudoporous (e.g. Fig. 178). These typically have a less-
regular arrangement of pseudopores than is found in frontal
walls. Non-pseudoporous terminal diaphragms may be planar
Or possess a central depression with pore (e.g. Fig. 238),
reminiscent of the ‘calcified terminal-vestibular membranes’
known from other cyclostomes (Boardman & McKinney,
1976; Boardman et al., 1992). In Meliceritites lorieri (Mich-
elin) some zooids possess two or even three pores in the
central depression. A third type of terminal diaphragm
comprises an apparently simple, flat plate (e.g. Fig. 252). The
exact location within the zooid of terminal diaphragms varies:
some are positioned at about the same level as the opercu-
lum, but others are situated more proximally, occasionally
beneath the apertural shelf (Fig. 252). Terminal diaphragms
in relatively proximal positions may underlie in-situ opercula,
as can be observed in thin sections or externally when in-situ
opercula are damaged (Fig. 240). This implies that opercula
do not necessarily have to be lost or shed before a terminal
diaphragm can be secreted.

Calcified opercula (Fig. 12) are known or inferred to be a
feature of all species of melicerititids. In most species they are
commonly found in-situ, but in a few species in-situ opercula
are extremely rare (e.g. Meliceritites transversa Canu &

Bassler). Opercula are occasionally found very close to the
growing edge of the colony, sometimes within incompletely-
formed apertures. Melicerititid opercula are undoubtedly
exterior walls (i.e. with cuticle-covered external surfaces
during life), even in the free-walled genera Foricula and
Biforicula. However, their microstructure and ultrastructure
differs from that of exterior frontal walls. The outer surface
of exceptionally well-preserved opercula is marked by a series
of radial fissures which are orientated parallel to local growth
direction (e.g. Fig. 88). Poorly-preserved opercula often have
a ‘recrystallized’ appearance with a strong fabric in the same
orientation as these fissures (e.g. Fig. 101). Pseudopores are
seemingly ubiquitous in melicerititid opercula but in only a
few species are they near circular in shape and approximately
evenly distributed over the surface of the operculum (e.g.
Fig. 214). More often they are arranged in a crescent parallel
to the distolateral edge of the structure and are slit-shaped,
elongated parallel to local growth direction, (e.g. Figs 69, 88,
140, 186, 254). The inner surfaces of opercula possess a pair
of ridges just inward of their lateral margins (Fig. 92). These
ridges, termed sclerites by analogy with similar thickenings
found in the opercula of cheilostomes, can be observed in
sections of in-situ opercula, opercula dissected out of aper-
tures, and are also sometimes visible as moulds left in the
sediment filling the zooidal chamber after loss of the opercu-
lum (Figs 176, 266). Melicerititid sclerites are generally most
pronounced close to the hinge line and become reduced
distally. They often slope inwards towards the median line of
the operculum. When the opercular pseudopores are
arranged in a crescent, they are found to open immediately
inwards of the sclerites. A shallow, dimple-like depression
can often be seen on the outer surfaces of well-preserved
opercula in the middle of the proximal edge, i.e. adjacent to
the median bar of the hingeline in articulated opercula.
External surfaces of opercula tend to lack ornament,
although slight patterning is found in a few species (e.g. Fig.
58).

Autozooids with an additional rim within the main aper-
ture are common in melicerititids (Figs 34, 45, 52-53, 55, 70,
89, 177) and were called regenerations by Levinsen (1912).
This phenomenon should not however be confused with
polypide regeneration, the routine process of formation of a
new polypide (essentially gut and tentacles) by a zooid in
which the old polypide has degenerated. Instead, analogy
with cheilostomes (see Banta, 1969; Taylor, 1988) suggests
that the ‘regenerations’ of melicerititids probably resulted
from the budding of a new zooid within the skeletal chamber
of a dead zooid and are better termed intramural buds. While
autozooids are often budded intramurally into the chambers
of old autozooids, eleozooids can also be intramurally bud-
ded into a host autozooidal chamber (Figs 19, 57, 111, 114,
156, 165, 171, 190, 195-196). Melicerititids are outstanding
among cyclostomes in their high frequencies of intramural
budding; Levinsen (1912) was able to point to only a few
indistinct cases of ‘regenerations’ in Entalophora madrepora-
cea and Hornera lichenoides among non-melicerititid cyclos-
tomes.

Eleozooids

Melicerititid eleozooids are the morphological analogues of
cheilostome avicularia in representing zooids with modified
opercula (and apertures). However, whereas avicularia
invariably have hypertrophied opercula, the opercula of

apertural shelf

hinge tooth

APERTURE

pseudopore

OPERCULUM (EXTERNAL)

sclerite

OPERCULUM (INTERNAL)

Fig. 12 Diagrammatic illustration of apertural and opercular
morphology in melicerititid cyclostomes.

eleozooids may be hypertrophied or of reduced size relative
to apertures of autozooids in the same colony. Eleozooids
have no close analogues among modern cyclostomes but
resemble the avicularia of cheilostomes. Avicularia are poly-
morphs with hypertrophied opercula, variously enlarged as
mandibles or hair-like setae, and probably with a range of
functions including defence, cleaning and even colony loco-
motion (see Winston, 1984, 1986, 1991). Normally, avicularia
are heterozooids which are unable to feed, but occasionally
they possess a feeding polypide and are therefore autozooidal
polymorphs (Silén, 1977; Cook, 1979). Several different
types of avicularia can be present within single colonies of
some cheilostome species. Similarly, melicerititids may have
several types of eleozooids within a single colony. There are
frequently two types of eleozooids, one of which is larger and
the other smaller than the autozooids in the colony. The
restricted apertures of many eleozooids leads to the inference
that they lacked a protrusible polypide, were incapable of
feeding and are therefore classified as heterozooids. How-
ever, some of the larger eleozooids have apertures of suffi-
cient size to have allowed protrusion of a polypide and it is
possible that they were autozooidal polymorphs capable of
feeding.

Three end-member categories of eleozooids can be recog-
nized, termed rostrozooids, trifoliozooids and demizooids
(Taylor, 1986a). Although most eleozooids are readily cat-
egorized into one of these types, some species possess eleo-
zooids which are consistently difficult to classify. For
example, the eleozooids of Meliceritites magnifica
(d’Orbigny) (see Taylor, 1986a: fig. 2D) are rostrozooid-like
in size but have trifoliate apertures, whereas the eleozooids of

P.D. TAYLOR

Reptomultelea reussi (Pergens) (see p. 94) have the size of
demizooids or trifoliozooids but apertures resembling rostro-
zooids. In addition, species with two types of eleozooids
occasionally possess zooids with morphologies intermediate
between the two types. In all three types of eleozooids, the
frontal walls of fixed-walled species have an equivalent
density of pseudopores as autozooids in the same colony. In
this respect eleozooids differ from gonozooids which are
characterized by a high density of pseudopores.

Rostrozooids (e.g. Figs 39, 48, 63, 142-145, 180, 201, 208,
223, 255, 295) have their apertures prolonged distally into a
shelf-like platform, the rostrum. The frontal area of a rostro-
zooid typically exceeds that of an autozooid in the same
colony, and the rostrum may occlude a distal zooid which
would otherwise have apparently developed into a normal
autozooid (i.e. the rostrozooid takes the position of two
autozooids on the colony surface; see Boardman in Board-
man et al., 1983: fig. 49.1 and 49.2). Zooids adjacent to the
rostrum may be effected by its presence, sometimes develop-
ing as kenozooids (Fig. 143) and sometimes as autozooids but
with obliquely-directed apertures (e.g., in Meliceritites lorieri
the apertures of autozooids on either side of the rostrum are
turned inwards towards the rostrozooid). Apertures of ros-
trozooids vary in morphology from almost semicircular and
only slightly larger than an autozooidal aperture, to long and
distally pointed, or long and distally rounded and with sides
parallel or distinctly spatulate. In-situ opercula appear to be
less common in rostrozooids than in autozooids, although this
has not been tested quantitatively. Like autozooidal oper-
cula, however, the opercula of rostrozooids often have a
cresentic arrangement of slit-shaped pseudopores. Inner sur-
faces have been observed in only a few cases. Surprisingly,
sclerites of the type seen in autozooidal opercula seem to be
lacking in rostrozooidal opercula of Meliceritites tuberosa
(d’Orbigny), although a marginal thickening does appear to
be present. Rostrozooids are seldom arranged in regular or
predictable patterns. In some species, however, rostrozooids
may be clustered in groups of two or three, and they often
occur in particularly high concentrations in the basal encrust-
ing parts of erect species. Whereas frontal walls are lacking in
the autozooids of the free-walled genera Foricula and
Biforicula, it is notable that a small area of pseudoporous
frontal wall is sometimes present in the rostrozooids.
Together with the gonozooids, which are also fixed-walled in
these genera, this gives colonies a mixed free-/fixed-walled
organization within zones of astogenetic repetition. Intramu-
ral buds are commonly found within rostrozooids. These can
be smaller rostrozooids (Figs 81, 145), tritoliozooids, demi-
zooids or autozooids (Fig. 242). Multiple intramural rostro-
zooids commonly occur in some species, often with the distal
ends of each successive rostrozooid being slightly elevated
relative to the previous rostrozooid so that the aperture
becomes inclined at a progressively greater angle to the
colony surface. Two demizooids may sometimes occur as
intramural buds within a single rostrozooid of Atagma.

Trifoliozooids (Figs 23, 164, 189) are distinguished by
having apertures with a trifoliate opening in the shape of an
inverted ‘TI’. On closer inspection of well-preserved
examples, the opening is usually seen to be set within a
D-shaped area which is occupied by the operculum on the few
occasions that this is observed in-situ. Apertures are smaller
than those of autozooids, sometimes only slightly so, how-
ever. The frontal wall of trifoliozooids bears the same density
of pseudopores as an autozooid and varies from a little

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 11

smaller to considerably smaller than that of an autozooid.
Like rostrozooids, trifoliozooids may be concentrated in the
encrusting bases of erect colonies. They may also occur with
increased frequency in disrupted areas where crowding
appears to have prevented the development of autozooids.
Some species of Meliceritella have reverse (dorsal) branch
surfaces composed of trifoliozooids, generally in combination
with kenozooids. Trifoliozooids frequently occur as intramu-
ral buds within autozooids and sometimes within rostrozoo-
ids.

Demizooids are small eleozooids with simple D-shaped
apertures and similarly shaped opercula. They show similari-
ties to trifoliozooids in size, variation and distribution. Spe-
cies of Afagma are particularly characterized by large
numbers of demizooids which typically outnumber and may
completely enclose the autozooids in the colony. Like trifo-
liozooids, demizooids can be found as intramural buds within
both autozooids and rostrozooids.

Kenozooids

In fixed-walled melicerititids kenozooids are generally sealed
by a calcified frontal wall, lack an aperture, and are identical
in morphology to the kenozooids found in other tubuliporine
cyclostomes. They are invariably smaller than autozooids and
often more irregular in outline shape. Kenozooids are typi-
cally found in areas of growth disruption, including anasto-
moses (Figs. 228-229), bifurcations (Fig. 68), adjacent to
gonozooids (Fig. 75) and eleozooids (Figs 143, 217), and also
in the encrusting bases of erect colonies (Fig. 32). Some
frontally-budded overgrowths commence with an apparent
kenozooid and may include other kenozooids within the
associated secondary zone of astogenetic change. Over-
growths of apparently open kenozooids, covering autozooidal
frontal walls but leaving their apertures uncovered, are
occasionally found in fixed-walled species (Fig. 79).

The small openings between autozooidal apertures of free-
walled melicerititids were considered by Taylor (1986a) to be
‘interzooidal spaces’ rather than kenozooids because of their
origin very close to the colony surface. These structures
resemble the cancelli of hornerid cyclostomes and are accord-
ingly given the same name. In neither melicerititids nor
hornerids is the phylogeny of the group sufficiently well-
known to determine whether the cancelli had an evolutionary
origin as modified zooids or are non-zooidal in origin.

Gonozooids

Gonozooids, like those of many species of cyclostomes, are
comparatively uncommon and have yet to be discovered in
some melicerititid species. When known they are very similar
to the gonozooids found in other tubuliporine cyclostomes,
and also in articulate cyclostomes. Variation of gonozooid
morphology within the melicerititids is slight. The distal
frontal wall is bulbous, bears a high density of pseudopores,
and in outline shape is usually longitudinally elliptical (e.g.
Figs 24-25, 41, 65, 71, 78, 86, 262), occasionally more
equidimensional (Fig. 236), and sometimes rounded-
subtriangular with an almost straight distal edge (Fig. 209).
Seldom do the neighbouring autozooids significantly indent
the outline of the distal frontal wall (cf. Fig. 209), and they
have never been found to pierce the frontal wall (cf. tubulipo-
rines such as Plagioecia). Asymmetrical distal frontal walls
are occasionally found in which the axis of the gonozooid

bends significantly from local colony growth direction and the
ooeciopore opens laterally or obliquely.

The proximal end of the gonozooid appear identical to an
autozooid — in fixed-walled species, it has a typically hexago-
nal frontal wall with the same density of pseudopores as an
autozooid and a D-shaped opening similar to an autozooidal
aperture. However, instead of being closed by an operculum,
the D-shaped opening ( ‘maternal aperture’) forms the point
of origin of the densely-pseudoporous distal frontal wall of
the gonozooid (Fig. 42). Sometimes the distal frontal wall is
initially a parallel-sided tube before dilation (Fig. 35), and in
a few species (e.g. Meliceritites dollfusi Pergens) it grows
proximally to cover the autozooid-like proximal frontal wall.

The ooeciopore is situated terminally and is usually trans-
versely elliptical (Figs 138, 153, 166, 222, 292), less often
subcircular (Fig. 65). Transverse width tends to be roughly
the same as that of autozooids in the same colony. Rarely, the
distal rim is prolonged into a slight tongue overhanging the
ooeciopore. Significant ooeciostomes are lacking, even in
well-preserved material where breakage can be ruled out.
Teratological specimens include partially-formed, aborted
gonozooids (Fig. 43), and coalesced gonozooids sharing a
single ooeciopore (Fig. 147)

In all melicerititids with suitably broken gonozooids, a low
ring diaphragm occurs in the distal part of the gonozooid just
proximal to the ooeciopore (Figs 25, 71). This structure was
first observed by Levinsen (1912) who termed it the atrial
ring and noted the presence of a similar structure in the
articulate cyclostome Crisia eburnea (Levinsen, 1912: pl. 7,
fig. 12).

Removal of the roof allows the floor of the gonozooid to be
examined. Traces of zooids which failed to reach the colony
surface are visible through the basal wall of the gonozooid
(e.g. Figs 24-25). The floor of the gonozooid may consist of a
series of walls occluding the underlying zooids (e.g. Taylor,
1986a: fig. 3B). These overgrown zooids usually lack frontal
walls, but in some cases possess a frontal wall and fully-
formed D-shaped aperture (e.g. Levinsen, 1912: pl. 7, fig.
13), a condition found especially in zooids located more
proximally beneath the dilated part of the gonozooid. Such
features seem to indicate that development of the dilated part
of the gonozooid was often retarded relative to the develop-
ment of nearby autozooids.

Intramural buds have never been seen within gonozooids,
nor have gonozooids been observed to originate as intramural
buds.

STRATIGRAPHICAL DISTRIBUTION

The 36 species of fixed-walled lamellar melicerititids
described here range from Albian to Campanian in age.
Although species belonging to other melicerititid genera
occur in the Barremian, Aptian, Maastrichtian and Palae-
ocene, there are no known species of Elea, Semielea or
Reptomultelea in deposits of these ages. The peak diversity
for lamellar melicerititid species occurs in the Cenomanian,
which contains 17 species. Diversities of 7-9 species charac-
terize the Turonian, Coniacian and Santonian, whereas the
Albian and Campanian contain only 3 and 2 species respec-
tively. This diversity pattern can be contrasted with the
pattern for melicerititid species as a whole (Taylor 1986a, fig.

4: nb. constructed from a less complete database), which
shows a Santonian peak in diversity. The anomolously large
number of lamellar species in the Cenomanian probably
correlates with the greater availability of nearshore localities,
where species of Reptomultelea are particularly common.

Stratigraphical ranges of species of Elea and Reptomultelea
are given in Figures 16 and 127 respectively. Most species are
recorded from one stratigraphical stage only, but a few range
through two or three stages. Better sampling will inevitably
extend these ranges and caution should therefore be exer-
cised when using melicerititids as age indicators.

SYSTEMATIC PALAEONTOLOGY

Specimen repositories and abbreviations. Specimens studied
are housed in the following collections: BGS, British Geo-
logical Survey, Keyworth; BMNH, The Natural History
Museum, London; DM, Dresden Museum (Reuss Collec-
tion); EM, Essener Museum; MNHN, Muséum National
d’Histoire Naturelle, Paris; PSUB, Goldfuss Collection, Uni-
versitat Bonn; SMD, Staatl. Museum Mineralogie und
Geologie, Dresden; USNM, National Museum of Natural
History, Smithsonian Institution, Washington; VH, Voigt
Collection, Geologisch-Paldontologisches Institut und
Museum, Universitat Hamburg; ZMC, Zoologisk Museum,
Copenhagen.

Methods of study. Almost all of the species described have
been studied primarily using scanning electron microscopy
(SEM), and whenever possible type specimens have been
scanned. Application of SEM is becoming increasingly essen-
tial in bryozoology, both for the precise characterization of
species and for their clear photographic illustration (Taylor,
1990). In the case of Semielea dichotoma, however, no
material was available for SEM. Most SEM has been under-
taken on uncoated specimens in an environmental chamber
(Taylor, 198665), using either ISI 60-A or ISI ABT-S55 scan-
ning electron microscopes. Unless otherwise stated, all of the
scanning electron micrographs depicted here are images
formed by back-scattered electrons (cf. secondary electron
images which are more conventional in SEM of coated
specimens, e.g. Fig. 180). It must be emphasized that the
magnifications of these figures are very approximate because
of highly imprecise machine readings.

Morphometrical determinations have been made using an
eyepiece micrometer affixed to a Wild M7 binocular micro-
scope. Time limitations have meant that for most species
autozooidal dimensions have been determined from a single
specimen only, preferably the holotype or lectotype. Intra-
colony variability within a species is therefore poorly known
and deserves future study. Ten autozooids per colony have
been measured in most species. Mean, standard deviation
(SD), coefficient of variation (CV) and observed range are
given. For eleozooids and gonozooids it has often been
necessary to take measurements from several colonies to
obtain a reasonable sample size. Figure 13 summarizes the
principal measurements made on autozooids (and eleozoo-
ids); see Pitt & Taylor (1990: fig. 2) for comparable measure-
ments made on gonozooids. When present in sufficient
numbers, apertural measurements were made from zooids
with in-situ opercula, as these have clearly-defined edges.

Orientated thin sections (longitudinal, tangential and trans-

P.D. TAYLOR

apertural
width

'
1
'
1
'
1

apertural
length

frontal
length

frontal
width

Fig. 13 Measured zooidal dimensions.

verse) were prepared for a small number of species using a
similar method to that described by Nye et al. (1972).

Order CYCLOSTOMATA Busk, 1852
Suborder TUBULIPORINA Milne Edwards, 1838
Family ELEIDAE d’Orbigny, 1852

REVISED DIAGNOSIS. Colony erect, dendroid, tubular, bifoli-
ate or fenestrate; or encrusting, multiserial and generally
multilamellar; overgrowths originating by intrazooecial fis-
sion and eruptive budding onto the colony surface, initiating
a secondary zone of astogenetic change. Skeletal organization
fixed-walled or free-walled, in some species varying accord-
ing to polymorph type. Interior wall microstructure tripartite,
apparently with a layer of transverse fibres flanked by lamel-
lar layers. Ancestrula with a short distal tube. Zooidal
polymorphism well-developed, most species possessing one
or more types of eleozooids in addition to the autozooids,
gonozooids and kenozooids. Autozooids fixed- or free-
walled, operculate; operculum calcified, articulated with the
straight proximal hingeline of the aperture, semicircular or
arch-like in outline shape and bearing sclerites on the inner
surface; peristome lacking. Eleozooids fixed- or free-walled,
with opercula elongated to form pointed, rounded or spatu-
late mandibles in rostrozooids, reduced relative to autozoo-
idal opercula in the smaller trifoliozooids and demizooids.
Gonozooids non-operculate, fixed-walled with distal frontal
walls densely pseudoporous and longitudinally elliptical
(occasionally subtriangular) in outline shape, neither pen-
etrated nor significantly indented by autozooids; ooeciopore
terminal, transversely elliptical to subcircular; ooeciostome
short; ring diaphragm (atrial ring) of low relief present just
proximal to ooeciopore. Intramural budding common, often
involving budding of a different polymorph type within the
skeletal chamber of a host zooid.

REMARKS. The Family Eleidae d’Orbigny, 1852 antedates
Melicerititidae Pergens, 1890 and, although priority is not
mandatory with regard to family names, Eleidae is here

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 13

preferred but the term ‘melicerititid’ is retained for vernacu-
lar purposes.

Melicerititids have a clear autapomorphy in the possession
of a calcified operculum, hinged to the proximal edge of the
aperture and able to seal the zooid on closure. No other
cyclostomes are known to have calcified (or uncalcified)
opercula. The high level of polymorphism in melicerititids
undoubtedly relates to the presence of this operculum which
can be modified to form various polymorphs, analagous to
the avicularia of cheilostomes derived by differentiation of
the operculum (Taylor, 19862).

Eight genera of melicerititids are recognized (Table 2).
These are distinguished principally according to the tradi-
tional character of colony-form. A full phylogenetic analysis
of the group will be necessary to determine whether zooid-
level characters may be significant in an improved generic
subdivision.

DISTRIBUTION. Barremian-Danian (or Montian), Europe
and western Asia.

Genus ELEA d’Orbigny, 1853

TYPE SPECIES. Bidiastopora lamellosa dOrbigny, 1850, des-
ignated by Gregory (1899, p. 299).

OTHER SPECIES. Elea elegantula sp. nov., Elea flabellata sp.
nov., Elea hexagona d’Orbigny, 1853, Eschara labyrinthica
Michelin, 1843, Elea mackinneyi sp. nov., Elea pseudolamel-
losa sp. nov., Elea subhexagona sp. nov., Eschara triangularis
Michelin, 1841, Elea viskovae sp. nov., Elea whiteleyi sp.
nov.

REVISED DIAGNOSIS. Eleid with bifoliate colony-form; auto-
zooids with a fixed-walled organization; cancelli lacking.

REMARKS. D’Orbigny’s (1853) original description of this
genus emphasized its similarities to Meliceritites from which it
was distinguished by the bifoliate form of the colony. Without
choosing a type species, d’Orbigny included the following ten
species in his new genus: Diastopora cervicornis Michelin,
1845, Bidiastopora ramossisima d’Orbigny, 1850, Eschara
ranvilliana Michelin, 1845, Elea calloviensis sp. nov., Elea
reticulata sp. nov., Eschara triangularis Michelin, 1841, Elea
rhomboidalis sp. nov., Elea turoniensis sp. nov., Elea lamel-

Table 2 Key to melicerititid genera.

1. Organization fixed-walled (i.e. autozooids with pseudoporous

frontal walls; cancelli lacking) ..............:.c:sccseeeeeeneeeeeene es 2
Organization free-walled (i.e. autozooids lacking
pseudoporous frontal walls; cancelli present) ..................+- 7
Za Colony entirely encrusting .-.-............-...0..00.- Reptomutltelea
CBO TY CEC SageccsadsaceacccededasqbouseccncocbcodosbsousbodoodsdasestGe 3
3. Branches bilaminate, strap-like or frondose ................. Elea
Branches dendroid, circular or subcircular in cross-section .. 4
Pememsraniches) HOMOW .2se-tecsts-s2-- ee sccssscse ce ospereueeadecsesss Semielea
EPG ANIGHESISOLIGUD «ap rderqaeystae ance ra dasigyde ere rewmcbise ote tceise ieee +elems 5
5. Autozooids absent on one side of branch .......... Meliceritella

Autozooids present around entire circumference of branch . 6
6. Autozooids surrounded by numerous small eleozooids with

IDES MEY OSEA DELTINIRES) orgococeccasducoecodnodaecoogncocuspsaseecod Atagma

THER WISE osha eed teeta ede ddasacs Meliceritites
7. Branches bilaminate, strap-like or frondose .......... Biforicula

Branches dendroid, circular or subcircular in

EROSS=SCCIl OM esate cia Aecichuaiiehontimagunten aemeasecdoeecgess Foricula

losa sp. nov., and Elea hexagona sp. nov. The first four
species are from the Jurassic and are not melicerititids; all
four were placed in synonymy with Multisparsa lamellosa
(Michelin, 1845) by Walter (1970). Elea reticulata from the
Neocomian is also not a melicerititid. Walter (1985) assigned
it to the tubuliporine genus Mesenteripora, as Mesenteripora
reticulata (d’Orbigny, 1853). E. turoniensis, from the Turo-
nian of Sainte-Maure (Indre-et-Loire), is represented by four
specimens registered as No. 6964 in the d’Orbigny Collection,
MNHRN, including one specimen in a glass tube here desig-
nated as the lectotype (Voigt photocard 6959). This too is not
a melicerititid. The remaining species listed by d’Orbigny are
true melicerititids and are correctly assigned to the Family
Eleidae.

Elea meridiana, described by Lang (in Woods, 1906: p.
283) from the Cretaceous [?Campanian] of Pondoland, South
Africa is a bifoliate tubuliporine, not a melicerititid. Three
figured syntype fragments are registered in the BMNH collec-
tions as D11834.

Among the eleven species validly assigned to Elea, a few
subgroupings may be recognized. One subgrouping com-
prises Ejea elegantula sp. nov. from the Lower Cenomanian,
E. subhexagona sp. nov. from the Upper Cenomanian and
Turonian, and E. hexagona d’Orbigny from the Santonian,
three species with adeoniform colonies and very similar
zooidal morphologies. They can be reasonably interpreted as
members of a clade exhibiting an evolutionary trend towards
an increase in the size of the autozooidal aperture relative to
the frontal wall. E. lamellosa (d’Orbigny) and E. pseudola-
mellosa sp. nov. can be distinguished only by the morphology
of the eleozooidal apertures which have an inverted T-shape
in the former (trifoliozooid-type) but are D-shaped in the
latter (demizooid-type). Other species of Elea (e.g. E. trian-
gularis (Michelin) and E. whiteleyi sp. nov.) are very distinc-
tive and more difficult to relate to congeneric species. E.
whiteleyi from the Lower-Middle Cenomanian may be a
member of the stem-group of Biforicula, judging by the
similarity of its small eleozooids to those of the earliest
species of Biforicula, B. multicincta from the Upper Cenoma-
nian (see Voigt 1989). Table 3 is a key to the identification of
species of Elea and should be used in conjunction with
Figures 14 and 15, which show the characteristic outline
shapes of the autozooidal and eleozooidal apertures respec-
tively.

It is worthwhile speculating on the phylogenetic origin of
Elea. Assuming that the genus originated from a fixed-walled
ancestor, there are two main possible ancestral genera:
Meliceritites and Reptomultelea. Origination from Meliceri-
tites would require a change in the mode of erect growth from
dendroid to bifoliate, whereas origination from Reptomulte-
lea would require the acquisition of erect growth. The latter
hypothesis is favoured for two reasons: (1) extensive
Reptomultelea-like bases are often present in species of Elea,
and (2) similarities exist between species of Elea and Repto-
multelea in the shape of the acuminate eleozooids (rostrozoo-
ids). Full phylogenetic analysis is, however, needed.
Stratigraphical distribution is not helpful in this respect as
Reptomultelea and Elea both have their earliest known occur-
rences in the Lower Albian while Meliceritites ranges back to
the Upper Barremian.

DISTRIBUTION. Lower Albian-Upper Campanian (Fig. 16) of
France, Germany, England and Kazakhstan.

ezace

E. labyrinthica _ E. whiteleyi E. elegantula

Ghose

E. subhexagona E. viskovae

E. hexagona

P. D. TAYLOR

e225

E. mackinneyi E. pseudolamellosa E. lamellosa
oe ii C
E. flabellata E.triangularis _S. vieilbanci

Fig. 14 Outlines of autozooidal aperture shapes in species of Elea and Semielea. Individual apertures were traced from SEM micrographs
and scaled using the mean apertural length determined for the species. Species are arranged according to apertural length.

Elea lamellosa (d’Orbigny, 1850) Figs 17-28

1850 Bidiastopora lamellosa d’Orbigny: 266.

1851 Bidiastopora lamellosa d’ Orbigny; d’Orbigny, pl. 625,
figs 11-15.

1853 Elea lamellosa (d’Orbigny); d’Orbigny: 632.

1853 Semielea plana d’Orbigny: 638, pl. 738, figs 12-14.

21853 Reptelea pulchella d’Orbigny: 642, pl. 738, figs 16-17.

1853. Semimutltelea irregularis d’Orbigny: 652, pl. 741, figs
6-8.

1853 Semimultelea gradata d’Orbigny: 653 (partim), ?non
pl. 741, figs 9-13.

1890 Semielea plana d’Orbigny; Pergens: 393.

1890 Elea lamellosa (d’Orbigny); Pergens: 398.

1899 Reptelea pulchella d’Orbigny; Gregory: 292, non fig.
Sl.

1899 Elea lamellosa (d’Orbigny); Gregory: 299.

non 1899 Semimultelea irregularis d’Orbigny; Gregory: 296,

fig. 32.

1912 Meliceritites plana (d’Orbigny); Levinsen: 43, pl. 5,
figs 11-12.

1912 Meliceritites lamellosa (d’Orbigny); Levinsen: 45, pl.
3, figs 1-9.

1985a_ lea lamellosa (d’Orbigny); Taylor, fig. 2F and G.
1985b Semielea plana d’Orbigny; Voigt: 631, pl. 3, figs
16-17.

MATERIAL. Type: no type specimens have been designated;
the d’Orbigny Collection, MNHN includes 8 glass tubes, all
registered as 8191, each of which contains specimens of Elea
lamellosa (e.g.Fig. 17). Some of these are from the two
localities given in the original species description (i.e. Tours
and Saintes); a lectotype could be selected from among these
topotypic syntypes but the identity of the species is not in
doubt.

Other material: MNHN d’Orbigny Colln 8191 (Voigt pho-
tocard 8310), Senonian [Santonian], Vendéme, France.
MNHN d’Orbigny Colln 8195, specimen labelled by E. Voigt
as the type of Semielea plana d’Orbigny (Voigt photocard
8319; figd by Voigt, 1985b: pl. 3, fig. 17), herein designated
the lectotype of S. plana, Senonian, ?Tours, France. MNHN
8201 d’Orbigny Colln, specimen labelled by E. Voigt as the
type of Semimultelea irregularis d’Orbigny (Voigt photocard
5733; figd by d’Orbigny, 1853: pl. 741, figs 6-8), herein
designated the lectotype of S. irregularis, Senonian, Tours,
France. BMNH D11786, Turonian, Touvent, Charente

viskovae elegantula labyrinthica subhexagona

0.1mm

flabellata hexagona whiteleyi

Fig. 15 Outlines of eleozooidal aperture shapes in species of Elea.
Individual apertures were traced from SEM micrographs and
scaled using the mean apertural length determined for the species.
Species are arranged according to apertural length.

Inferieure, Jukes Browne Colln; D54296, Coniacian, Tours,
Indre-et-Loire, Voigt Colln; D59166-7, Coniacian, Craie de
Villedieu, Rue St Barthelemy, Tours, Taylor and Hammond
Colln; D36195, Coniacian, St Paterne, Indre-et-Loire, Per-
gens Colln; D36176, Senonian, St Antoine-du-Rocher, Indre-
et-Loire, Pergens Colln; D36060-2, Senonian, Vendome,
Loir-et-Cher, Pergens Colln; D53637, D59159, Coniacian or
Santonian, Craie de Villedieu, Villedieu, Loir-et-Cher, Gale
Colln; D59165, Coniacian or Santonian, Craie de Villedieu,
Chateau Mbr, between Villedieu and Trehet on D80, Taylor
and Hammond Colln; D58849 (sample), Santonian, Craie de
Villedieu, Chateau Mbr, Bed 17 of Jarvis et al. (1982),
Nowicki Colln; D58843 (sample), D59160-2, D59163
(sample), D59164, D59202-3, Santonian, Craie de Villedieu,
Bouchardiére Mbr, Bed 20 of Jarvis et al. (1982), Villedieu,
Gale Colln; D58839 (sample), Santonian, Craie de Villedieu,
Bouchardiére Mbr, Bed 22a of Jarvis et al. (1982), Villedieu;

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

D59168 (sample), Santonian, Craie de Blois, Rue St Bar-
thelemy, Tours, Taylor and Hammond Colln; D59169-72,
Santonian, Craie de Saintes Fm., Voiville Mbr, les Arcivaux-
Portublé, Saintes, France. USNM 2737-1 to 4, Coniacian,
Villedieu (sections including duplicate acetate peels in
BMNH). ZMC M40, [?Coniacian or Santonian], St Antoine
du Rocher, France. ZMC Levinsen material unnumbered,
Coniacian, Villedieu, Canu Colln.

Un-numbered VH material from the following localities:
Coniacian, Tours; Coniacian, Villedieu; Coniacian, St Chris-
tophe, Indre-et-Loire; Santonian, Vendome; Santonian,
Merpins, Charente-Maritime; Coniacian, Joué-les-Tours,
Indre-et-Loire (labelled Semimutltelea irregularis).

DESCRIPTION. Colony normally bifoliate, foliaceous (Fig.
17), with broad, folded fronds (Fig. 18), about 0-6-0-8 mm
deep, anastomosing and giving rise to daughter fronds per-
pendicular to their surfaces; overall shapes of complete
colonies are flattened spheroids, up to at least 40 mm in
diameter (e.g. BMNH DS59159); parts of some colonies,
including extensive colony bases, are unilamellar with a
concentrically ridged exterior wall forming the basal side.
Growing edges revealing several generations of buds, often
occluded by a combination of small eleozooids and kenozoo-
ids. Overgrowths common, formed by eruptive budding onto

Table 3 Key to the species of Elea. Note that because this key
places a high reliance on eleozooids, which may not be developed
in every specimen, identifications should be carefully checked
against the full descriptions; furthermore, it is possible that new
material will reveal the presence of eleozooids in species in which
they are currently unknown.

1. Autozooidal aperture more than 0-25 mm long, bell-shaped;
eleozooids unknown ...................0665 Elea triangularis (p.34)

Autozooidal aperture less than 0-25 mm long ................... 2
2. Eleozooids with long rostra (rostrozooids) present ............ 3
ROSTLOZOOIGSHACKIN GY ater awocecdereceirertsearestelseciaatetecsnaesen 10

3. Small eleozooids with D-shaped apertures (demizooids)
AOU ANLMeREee Meee tei cc aecececeescnescocea cous Elea whiteleyi (p.39)

WETUIZOOIGSHACKIN Pale sencnce et escees see ashe re cern esieesecceesteneneee 4
4. A single large and prominent barkhan dune-shaped tubercle

situated distally of the zooidal apertures ..... Elea labyrinthica

(p.25)

AO CISC MP Ee Pec tee ose ce autie cnsiene etGau ceiceiteipateisnicl vadeneee active cles 5

5. Reduced frontal wall occupying less than half of the
autozooidal frontal surface; tuberculate zooidal

OU ATES rece eciar .ctiescctecdewe cits ceetas Elea flabellata (p.20)
(CHRGIWHISS sendeactnesdassndteonosee dead sbanecnaeccno nBEenede Sa pascee recor 6
6. Eleozooids with broad, well-rounded rostra ..................... V/
Eleozooids with narrow, pointed rostra ..................002eeee 8

7. Autozooids with broad apertural shelves; rostrozooid apertures
less than 1-5 X longer than wide ........... Elea viskovae (p.38)
Autozooids with apertural shelf lacking or very narrow;
rostrozooid apertures about twice as long as wide
_ coc a0ggU dR OSAP EEBREEB SEE BU ABEAREBRDAERERr EEE Elea mackinneyi (p.29)

8. Autozooids with a tubercle distal of the aperture

De ta steam ees aaa ales Saisie wlesbisive sels elises Elea hexagona (p.23)
Tubercle lacking

9. Autozooid frontal width less than 0-25 mm
copvaaatapnacete SaReCeEe cece cose acne Eee er eecernnetts Elea elegantula (p.19)
Autozooid frontal width more than 0-25 mm
"Eg :O GEE RDC ODE Re Done CEE EEREEEe Rec Cor nc nent Elea subhexagona (p.32)

10. Eleozooids with inverted T-shaped apertures
(GitfaltoZ0O1dS) pes sceeceeeneteeesssewencie cre Elea lamellosa (p.14)
Eleozooids with D-shaped apertures (demizooids)
oti PCOS ED SEROUS BUEN ARLE ec Oe Elea pseudolamellosa (p.30)

NVIDSVINOD
NVINOLNVS
NVINVdWYVD

NVINVWON39
NVINOYNL

triangularis

labyrinthica

elegantula
mackinneyi
whiteleyi
subhexagona
viskovae
lamellosa
flabellata
hexagona

pseudolamellosa

Fig. 17 Elea lamellosa (d’Orbigny, 1850), MNHN d’Orbigny
Collection 8191 (Voigt photocard 8310), Senonian [Santonian],
Vendome, France; photograph of frond fragment, x 4-8.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 17

the surface of a frond, initiating a zone of secondary astoge-
netic change, becoming subcircular in outline; pseudoances-
trula an autozooid; autozooids in zone of change in old
overgrowths often with intramurally budded eleozooids. Zoo-
ids generally arranged in quincunx; organization fixed-
walled.

Autozooids (Fig. 20) medium-sized, frontally hexagonal or
diamond-shaped, about 1-5 x longer than wide, outline
~ extended distally and rounded by apertural rim; frontal wall
flat or very slightly convex, with circular pseudopores; zoo-
ecial boundaries distinct, raised. Apertures (Fig. 22) small,
longitudinally elongate, 1-1—-1-4 x longer than wide, attaining
maximum width at or a little distally of the hinge line,
well-rounded distally; apertural shelf narrow; apertural rim
prominent, moderately broad, continuous with raised wall
forming remainder of zooecial boundary; hinge line with
median bar. Opercula (Fig. 21) very often preserved in-situ,
prominent, surface convex, pseudopores not observed. Ter-
minal diaphragms not observed. Intramural eleozooids (Fig.
19) common, their inverted T-shaped apertures slightly
smaller than those of primary eleozooids and occupying the
distal half of the host autozooid aperture, a slightly concave
pseudoporous exterior wall fills the proximal part of the host
autozooid aperture.

Eleozooids abundant, distributed widely across fronds,
sometimes clustered in small groups (Fig. 19), frontally 1-5-2
x longer than wide, typically a little narrower and more
pointed distally than autozooids but of about the same length;
frontal wall with circular pseudopores in the same density as
those of the autozooids. Aperture (Fig. 23) small, longitudi-
nally elongate, about 1-5-2 x longer than wide, attaining
maximum width at the hinge line, inverted T-shaped in
outline through indentation of lateral margins by the rostral
shelf. Opercula rarely preserved in-situ, longitudinally elon-
gate, rounded distally. Intramurally budded eleozooids
within host eleozooids not observed.

Kenozooids often present in association with eleozooids,
generally with a smaller frontal area than other zooidal
polymorphs.

Gonozooids (Figs 24-25) common, sometimes paired,
large, longitudinally elongate; frontal wall with a short
parallel-sided portion emerging from the maternal zooidal
aperture, dilating into an ovoidal bulbous portion almost
twice as long as wide. Ooeciopore (Fig. 24) transversely
elongate, 2-2-5 x wider than long, in outline often kidney-
shaped due to the presence of an indenting proximal
hemiseptum. Atrial ring (Fig. 25) well-developed.

Thin sections show median budding lamina (Figs 26-27)
straight to slightly sinuous, similar to the normal interzooidal
walls in microstructure; zooecia up to at least 0-63 mm in total
length; frontal walls thinning distally towards hinge line;
opercula (Fig. 28) thin, often less than 0-015 mm in thickness,
sclerites present; interzooidal walls about 0-015 mm thick
near centre of branch, increasing to 0-03-0:05 mm near
surface; basal diaphragms occasionally present in proximal
parts of zooids.

Fig. 24 Elea lamellosa (d’Orbigny, 1850), BMNH D59203,
Santonian, Craie de Villedieu, Bouchardiére Member, Villedieu,
France; de-roofed gonozooid with intact ooeciopore, x 45.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from BMNH
D59164)
frontal length: mean = 0-49 mm; SD = 0-040 mm;
CV = 8:2; range = 0-45-0-57 mm
mean = 0-32 mm; SD = 0-015 mm;
CV = 4-9; range = 0-30-0-35 mm
mean = 0-19 mm; SD = 0-008 mm;
CV = 4-2; range = 0-18-0-20 mm
mean = 0:16 mm; SD = 0-013 mm;
CV = 8-1; range = 0-14-0-18 mm

eleozooids (10 zooids from BMNH D59164)

frontal length: mean = 0-50 mm; SD = 0-054 mm;
CV = 10-9; range = 0:42-0:59 mm
mean = 0-28 mm; SD = 0-046 mm;
CV = 16-7; range = 0-18-0-33 mm
mean = 0-10 mm; SD = 0-008 mm;
CV = 8-1; range = 0-09-0-11 mm

frontal width:
apertural length:

apertural width:

frontal width:

apertural length:

Figs 18-23 Elea lamellosa (d’Orbigny, 1850). 18, BMNH D53637, Coniacian or Santonian, Craie de Villedieu, Villedieu, Loir-et-Cher,
France, frond fragment with two broken gonozooids and a partly enveloped, fouling oyster, x 7-5. 19, BMNH D59164, Santonian, Craie
de Villedieu, Bouchardiére Mbr, Villedieu, primary eleozooids, operculate autozooids and a damaged intramural eleozooid (lower
right), < 75. 20-21, ZMC Levinsen Collection M40, [?Coniacian or Santonian], St Antoine du Rocher, France; 20, regularly-arranged,
operculate autozooids, x 74; 21, autozooidal operculum, x 220. 22-23, Santonian, Craie de Villedieu, Bouchardiére Member, Villedieu;
22, BMNH D59203, autozooidal aperture, x 285; 23, BMNH D59202, eleozooid aperture, x 330.

Fig. 25 Elea lamellosa (d’Orbigny, 1850), BMNH D59202,
Santonian, Craie de Villedieu, Bouchardiére Member Villedieu,
France; de-roofed gonozooid exposing entrance to maternal zooid
and atrial ring, x 45.

mean = 0-06 mm; SD = 0-009 mm;
CV = 16-6; range = 0-05-0-08 mm

gonozooids (6 zooids from sample BMNH D59163)
total frontal length: range = 2-22-2-63 mm
distal frontal wall
length:

frontal width:
ooeciopore length:
ooeciopore width:

apertural width:

range = 1:86-2:25 mm
range = 1-10-1-31 mm
range = 0-08—0-11 mm
range = 0-17-0-20 mm

REMARKS. Elea lamellosa is an extremely abundant bryo-
zoan in the Coniacian-Santonian calcarenites of the Loire
region, notably the Craie de Villedieu, and in approximately
contemporaneous deposits in the Aquitaine Basin, for
example at Saintes. It is the only bifoliate melicerititid to
occur commonly in these beds, in which dendroid forms such
as Meliceritites magnifica, M. ornata and M. tuberosa tend to
dominate (Fig. 3). The foliaceous shape (‘eschariform’) of the
colonies contrasts with most other bifoliate melicerititids
which have strap-like branches (‘adeoniform’). Although
present in several species of Meliceritites and also in Semielea
vieilbanci, no other species of Elea possess small eleozooids
with inverted T-shaped apertures. These trifoliozooids are
the principal means of distinguishing between E. lamellosa
and the closely similar but stratigraphically younger E.
pseudolamellosa (p. 30).

P.D. TAYLOR

Figs 26-28 Elea lamellosa (d’Orbigny, 1850), photomicrographs of
thin sections of specimens from the Coniacian [?Santonian] of
Villedieu, Loir-et-Cher, France. 26-27, USNM 2737-2; 26,
transverse section, x 26; 27, detail showing median budding
lamina and eleozooid (top centre), x 100. 28, USNM 2737-1,
longitudinal section of an autozooid with in-situ operculum, X 97.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 19

Semielea plana d Orbigny (see Voigt, 1985b: pl. 3, fig. 17) isa
unilamellar fragment of Elea lamellosa with the beginnings of an
eruptive overgrowth, and Semimultelea irregularis d’Orbigny is
the multilamellar base of an Elea lamellosa colony. The identity
of Reptelea pulchella is uncertain. The lectotype (herein desig-
nated) of R. pulchella is MNHN d’Orbigny Collection 8196, the
specimen in a tube labelled by E. Voigt as the type (Voigt
photocard 8316). This Senonian specimen consists of a small
colony encrusting a bivalve shell fragment. Eleozooids are
lacking. It may be a young stage of E. /amellosa or perhaps the
base of another melicerititid species.

Semimultelea gradata d’Orbigny is represented by two speci-
mens registered as 8202 in the d’Orbigny Collection, MNHN.
One of the specimens is from Tours and appears to be the basal
part of a colony of Elea lamellosa. The second specimen, from
Meudon (Campanian), labelled by E. Voigt as the type (Voigt
photocard 5747), is herein designated as the lectotype. This is
probably the base of an indeterminate erect melicerititid, and is
unlikely to be Elea lamellosa in view of the otherwise absence of
this species in the chalky facies of Meudon.

Semimutltelea irregularis d’Orbigny was selected as the type
species of Semimultelea d’Orbigny by Gregory (1899: p. 296).
Unfortunately, Gregory’s material of supposed S. irregularis
is a mixture of indeterminate Turonian melicerititids with
overgrowths, none of them conspecific with true S. irregularis
(= Elea lamellosa). Nevertheless, the type species selection
should stand, making the genus Semimultelea an objective
junior synonym of Elea.

DISTRIBUTION. ?Turonian, Coniacian-Santonian of the Loire
region and Aquitaine, France. The Turonian record is based
on an old specimen (BMNH D11786) and, especially in view
of correlation problems, should be regarded as doubtful.

Elea elegantula sp. nov. Figs 29-43

1985a_ Elea sp. Taylor, fig. 2H and I.

MATERIAL. Holotype: VH 10451, Lower Cenomanian,
Milheim/Ruhr, Germany. Paratypes: VH 9847-8, 10452,
10473, same horizon and locality as paratype. Other material:
numerous un-numbered VH topotype specimens.

NAME. With reference to its elegant appearance.

DESCRIPTION. Colony bifoliate with apparently short
branches, generally narrow basally and expanded distally,
ranging from about 0-7-4 mm in width but typically about 1-6
mm wide, and approximately 0-4 mm deep, often divided
distally and frequently twisted. Zooid arrangement varies
from regular quincunx to poorly-defined transverse rows,
zooids close to the branch margin slightly divergent; organi-
zation of all polymorphs fixed-walled. Overgrowths (Fig. 33)
may arise through eruptive budding onto branch surface;
pseudoancestrula operculate, frontal wall negligable; zone of
secondary astogenetic change characterized by autozooids
with smaller and more rounded apertures than those in zones
of astogenetic repetition. Colony base (Figs 37—38) extensive,
giving rise to multiple erect branches whose budding laminae
are orientated parallel to local orientation of basal zooids.
Ancestrula not observed.

Autozooids (Fig. 29) of moderate size, with pseudoporous
frontal walls occupying half or more of the frontal area, about
2 X longer than wide, generally elongate hexagonal in shape
with rounded distal borders; zooecial boundaries raised and
well-defined. Apertures (Fig. 30) small, longitudinally elon-

gate, up to 1-3 x longer than wide, attaining maximum width

a little distal of the hinge line, rounded distally; apertural
shelf well-developed, broadening distally; hinge line with a
median ridge; apertural rim prominent. Opercula (Fig. 34)
frequently preserved in-situ, surface moderately convex, a
flattened median proximal area often evident, bearing about
25 elongate pseudopores in a crescent close to the lateral/
distal edge. Autozooids and more commonly small eleozoo-
ids (Fig. 31) may be budded intramurally within autozooids.
Terminal diaphragms, located below level of apertural shelf,
very occasionally present.

Eleozooids (Figs 29, 39-40) abundant, most positioned
close to branch margins, especially in the recesses of branch
divisions, small examples often associated with occluded
growing edges. Frontal walls generally a little narrower than
those of autozooids, with a similar concentration of
pseudopores. Apertures variable in length, most are long,
about 3 x longer than wide, tapering distally with concave
lateral edges and a well-developed shelf-like rostrum (typical
rostrozooids), but these grade into others which are short and
subtriangular; maximum apertural width attained at hinge
line level. Opercula not observed. Intramural budding of
small eleozooids within large eleozooids (Fig. 40) is common,
the rostrum being infilled by pseudoporous exterior wall.

Kenozooids of varying size and shape common, mainly
located at occluded growing edges, in recesses of branch
divisions and in encrusting bases.

- Gonozooids (Figs 41-43) common, sometimes more than
one per branch fragment. Distal frontal wall emerging from
maternal aperture initially parallel-sided, often for a consid-
erable distance, before becoming densely pseudoporous,
bulbous and longitudinally ovoidal in shape. Ooeciopore
positioned beyond inflated part of frontal wall, transversely
elliptical, about one-and-a-half times as wide as long, an
internal hemiseptum indenting the proximal edge to give a
kidney-shaped deep outline. Atrial ring present in at least
some gonozooids.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from holotype VH
10451)

frontal length: mean = 0:46 mm; SD = 0-026 mm;
CV = 5-5; range = 0-42-0-51 mm
mean = 0:23 mm; SD = 0-008 mm;
CV = 3-4; range = 0-23-0-24 mm
mean = 0-17 mm; SD = 0-010 mm;
CV = 6-1; range = 0:17-0:20 mm
mean = 0-15 mm; SD = 0-009 mm;
CV = 6:5; range = 0-14-0-17 mm
eleozooids (10 zooids from holotype VH 10451)

frontal length: mean = 0-60 mm; SD = 0-139 mm;
CV = 23-1; range = 0-32-0-74 mm
mean = 0-20 mm; SD = 0-020 mm;
CV = 10-0; range = 0-17-0-23 mm
mean = 0-32 mm; SD = 0-092 mm;
CV = 29-3; range = 0-12-0-41 mm
mean = 0-09 mm; SD = 0-012 mm;
CV = 12:7; range = 0-08-0-11 mm
gonozooids (10 zooids from holotype VH 10451

and un-numbered VH specimens)

frontal length: mean = 1:69 mm; SD = 0-227 mm;
CV = 13-4; range = 1-29-2-03 mm
mean = 1:06 mm; SD = 0-162 mm;
CV = 15-2; range = 0-74-1-31 mm

frontal width:
apertural length:

apertural width:

frontal width:
apertural length:

apertural width:

dilated frontal
length*:

P.D. TAYLOR

Figs 29-32 Elea elegantula sp. nov., Lower Cenomanian, Milheim/Ruhr, Germany. 29-30, VH 10451, holotype; 29, autozooids (some with
opercula) and eleozooids, x 60; 30, autozooidal aperture, x 250. 31, VH 10452, small intramural eleozooid with partially preserved
operculum, x 250. 32, VH 10473, autozooids, small primary eleozooids, intramural eleozooids and kenozooids in encrusting colony
base, x 42.

mean = 0-83 mm; SD = 0-134 mm;
CV = 16-2; range = 0-63-1-01 mm
mean = 0:08 mm; SD = 0-011 mm;
CV = 12-8; range = 0-06-0-09 mm
mean = 0:14 mm; SD = 0-019 mm;
CV = 13-2; range = 0:12-0:18 mm
(*1.e. that part of the frontal wall beyond the narrow, tubular
proximal frontal wall)

frontal wall width:
ooeciopore length:

ooeciopore width:

REMARKS. Particularly characteristic of this species are the
long and narrow eleozooidal apertures and the thick autozoo-
idal apertural rims. E. elegantula resembles E. hexagona
d’Orbigny, 1853, from the Santonian but lacks the tubercle
developed distally of the autozooidal aperture, and has
frontal walls which occupy a larger proportion of the autozoo-
idal frontal area. It is also very similar to the late
Cenomanian/Turonian species E. subhexagona sp. nov. but
has somewhat smaller zooids with relatively more extensive
frontal walls. E. mackinneyi sp. nov., which occurs with E.
elegantula at Miilheim/Ruhr, differs in having broad eleozoo-
idal apertures and more rounded autozooidal apertures.

DISTRIBUTION. Lower Cenomanian of Mtlheim/Ruhr, Ger-
many.

Elea flabellata sp. nov. Figs 44-48

MATERIAL. Holotype: VH 10448, Lower Santonian, Grube
Lengede-Broistedt, near Braunschweig, Germany. Other
material: VH unnumbered, 3 abraded fragments from the
same horizon and locality probably belong to this species.

NAME. Flabellum, fan (L.), with reference to the flabellate
shape of the holotype colony.

DESCRIPTION. Colony bifoliate, consisting in the holotype of
a single non-bifurcating, flabellate branch, 3-2 mm wide
proximally, expanding to a width of 14 mm distally, about 1-5
mm thick, distinctly elliptical in cross-section at the proximal
fracture. Autozooids arranged in approximate quincunx;
organization fixed-walled. Colony base and overgrowths not
observed.

Autozooids (Figs 44-45) of small size, frontally a little less

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 21

Figs 33-36 Elea elegantula sp. nov., Lower Cenomanian, Mulheim/Ruhr, Germany. 33, VH 10452, overgrowth origin, x 85. 34, VH 9848,
operculate autozooid (left) and autozooid containing intramural autozooid (right), x 115. 35, VH 9847, proximal part of a gonozooid
showing tubular frontal wall emerging from an autozooid-like aperture, x 75. 36, VH 10451, ooeciopore, x 245.

than twice as long as wide and typically hexagonal in outline
shape; zooidal boundaries marked by a broad raised area of
calcification prolonged into low, blunt tubercles at the cor-
ners of zooids; frontal wall pseudoporous, occupying a very
small proportion of the frontal surface (Fig. 47), considerably
smaller than the autozooidal apertures. Apertures (Fig. 47)
medium-sized, slightly longitudinally elongate or equidimen-
sional, attaining maximum width at about mid-length, well-
rounded distally; apertural shelf narrow; hinge line with a
median bar. Opercula (Fig. 46) occasionally preserved
in-situ, surface convex; pseudopores radially elongate,
present over entire surface of operculum. Terminal dia-
phragms not observed. Intramurally budded autozooids (Fig.
45) infrequent, equipped with a thin, raised secondary aper-
tural rim.

Eleozooids (Fig. 48) abundant, scattered, not concentrated
at branch margins, frontally on average 1-8 x longer than
wide and about 1-5 x the size of an autozooid; frontal wall
pseudoporous, occupying a small proportion of the frontal
surface but larger than an autozooidal frontal wall. Aperture
moderately elongate, about 1-5 x longer than wide, widest at
the hinge line, tapering distally to form a narrow, parallel-

sided rostrum which is approximately half the total length of
the aperture, rounded distally. Opercula observed in-situ.
Intramurally budded autozooids possibly present but rare;
intramural eleozooids common, many with apertures facing
proximally in a plane oblique to that of the host aperture.

Gonozooids represented by a single example which is both
abraded and incompletely formed; longitudinally elongate in
outline; ooeciopore and atrial ring not observed.

REMARKS. Only a single adequately-preserved fragment is
known of this species. However, the specimen is sufficiently
different from other species of Elea to be confident that it
represents a new species. Although the eleozooids of E.
flabellata are similar to those present in several other species
of Elea (e.g. E. labyrinthica (Michelin) and E. hexagona
d’Orbigny), the thick tuberculate calcification which sur-
rounds these as well as the autozooids is very characteristic of
the species, as are the much reduced frontal walls. The
distribution of pseudopores over the entire surface of the
autozooidal operculum is a feature apparently unique to E.
flabellata among species of Elea, although it is present in
some species of Meliceritites (notably M. dollfusi Pergens)

22 P.D. TAYLOR

Figs 37, 38 Elea elegantula sp. nov., VH 10473, Lower Cenomanian, Miilheim/Ruhr, Germany; 37, colony base encrusting a shell, giving
rise to three erect branches (upper left and right) and forming an arch (lower centre) probably where a soft-bodied organism was
overgrown, X 17; 38, base of erect branch, x S0.

Figs 39-41 Elea elegantula sp. nov., VH 10451, Lower Cenomanian, Milheim/Ruhr, Germany; 39, large eleozooid, x 160; 40, large
eleozooid containing small intramural eleozooid, x 145; 41, gonozooid with broken roof, x 65.

and in Reptomultelea scanica sp. nov. E. flabellata bears a DISTRIBUTION. Lower Santonian of Broistedt, Germany.
resemblance to Biforicula nodulifera (Voigt), also from the

Santonian of West Germany. Abraded examples of these two MEASUREMENTS.

species could be difficult to tell apart were it not for the rather autozooids (10 zooids with in-situ opercula from holotype VH
more pointed apertures of B. nodulifera. 10448)

apertural width:

frontal width:

_ apertural width:

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

i)
WwW

Figs 42, 43 Elea elegantula sp. nov., Lower Cenomanian, Mtlheim/Ruhr, Germany 42, VH 10451, inflated frontal wall of gonozooid
emerging from an autozooid-like aperture, x 145. 43, VH 10452, gonozooid at the edge of a branch which was apparently aborted after
growing the proximal part of the inflated frontal wall; zooids which would have formed the floor of the gonozooid are sealed by exterior
walls, x 80.

mean = 0-41 mm; SD = 0-027 mm;
CV = 6-7; range = 0:36-0:45 mm
mean = 0-27 mm; SD = 0-015 mm;
CV = 5-8; range = 0-24-0-29 mm
mean = 0-21 mm; SD = 0-010 mm;
CV = 4-9; range = 0-20-0-23 mm
mean = 0-20 mm; SD = 0-007 mm;
CV = 3-6; range = 0-20-0-21 mm

frontal length:
frontal width:

apertural length:

- eleozooids (10 zooids from holotype VH 10448)

mean = 0-66 mm; SD = 0-048 mm;
CV = 7-3; range = 0:57-0:75 mm
mean = 0-36 mm; SD = 0-020 mm;
CV = 5-5; range = 0-35-0-39 mm
mean = 0-40 mm; SD = 0-042 mm;
CV = 10-5; range = 0-33-0-45 mm
mean = 0-27 mm; SD = 0-018 mm;
CV = 6-6; range = 0-26-0-:30 mm

frontal length:

apertural length:

gonozooid (abraded, incomplete zooid from holotype VH
10448)

total frontal length:
frontal width:

>1-73 mm
ca 1:28 mm

Elea hexagona d Orbigny, 1853 Figs 49-56

1853 Elea hexagona d’Orbigny: 633, pl. 738, figs 1-4.
1889 Elea hexagona d’Orbigny; Pergens: 398.

non 1897 6 Elea hexagona d’Orbigny; Canu: 753.

1899 Elea hexagona (d’Orbigny); Gregory: 303.

1912 Meliceritites hexagona (d’Orbigny); Levinsen: 43, pl.
5, figs 3-5.

MATERIAL. Lectotype (herein designated): MNHN
d’Orbigny Collection 8192 (Fig. 49), fragment in tube
labelled ‘Type’ by E. Voigt (Voigt photocard 5791), Senonian
[Santonian], Vendome, Loir-et-Cher, France. Paralectotype:
one of the two other fragments registered with lectotype as
MNHN 8192 (the third fragment is ?Foricula). Other mate-
rial: ZMC Levinsen Colln M38 and M39, Santonian, Ven-
déme, France; ZMC Levinsen Colln, about ten un-numbered
topotypes.

DESCRIPTION. Colony bifoliate with branches (Figs 49-50)
bifurcating, strap-like, narrow, about 1-7-2:3 mm wide.
Autozooids usually arranged in approximate quincunx; orga-
nization fixed-walled. Colony base not observed. Over-
growths produced by eruptive budding onto the branch
surface present.

Autozooids (Fig. 51) moderately large; frontal surface
usually hexagonal with edges parallel to long axis or
diamond-shaped, elongate, about twice as long as wide;
frontal wall small in area, with subcircular pseudopores;
zooecial boundaries formed mainly by apertural rims, raised.
Apertures (Fig. 52) medium-sized, longitudinally elongate,
about 1:2 x longer than wide, attaining maximum width

P.D. TAYLOR

Figs 44-47 Elea flabellata sp. nov., VH 10448, holotype, Lower Santonian, Grube Lengede-Broistedt, near Braunschweig, Germany; 44,
autozooids and three eleozooids, < 50; 45, autozooids, an eleozooid, and an intramural autozooid (top centre left), x 70; 46, autozooidal
operculum, Xx 200; 47, autozooidal aperture and reduced frontal wall, x 160.

between hinge line and mid-length, rounded distally; aper-
tural shelf variably developed; apertural rim moderately
prominent proximally, distally forming a conspicuous, deep
tubercle; hinge line bowed. Opercula (Fig. 56) often pre-
served in-situ, surface convex, pseudopores not evident in
poorly-preserved available material. Terminal diaphragms
not observed. Intramurally budded autozooids (Figs 52-53)
common, some having apertures in same plane as host
aperture but significantly smaller and more rounded distally,
others having apertures in an oblique plane facing proximally
relative to branch orientation and possessing a very promi-
nent tubercle distal to the aperture.

Eleozooids (Figs 51, 53-54) abundant, located particularly
at branch margins and in bifurcations, about 2-5-3 x longer
than wide, frontally slightly wider and significantly longer
than autozooids. Aperture elongate, about 2-2-5 x longer
than wide, attaining maximum width at the hinge line and
tapering distally to a long, narrow rostrum with a rounded
end. Opercula not observed in-situ.

Gonozooid unknown.

MEASUREMENTS (estimated from SEM micrographs).
autozooids

frontal wall length: ca 0:43-0:50 mm
frontal wall width: ca 0:22-0:26 mm
apertural length: ca 0-18-0-21 mm
apertural width: ca 0:16-0:18 mm

eleozooids

frontal wall length:
frontal wall width:
apertural length:
apertural width:

ca 0:62-0:83 mm
ca 0:25-0:27 mm
ca 0:36-0:50 mm
ca 0:17-0:23 mm

REMARKS. The supposed Elea hexagona described by Canu
(1897b) from the Cenomanian of Saint-Calais is here included
in E. subhexagona sp. nov. This species more closely
resembles E. hexagona than any other species of Elea but
lacks the very deep and prominent tubercle distal to the

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 25

Fig. 48 lea flabellata sp. nov., VH 10448, holotype, Lower
Santonian, Grube Lengede-Broistedt, near Braunschweig,
Germany; eleozooid, x 185.

autozooidal aperture, which is a characteristic feature of E.
hexagona. The larger size of the autozooidal apertures
enables distinction between E. hexagona and E. labyrinthica.
Filliozat (1908) records E. hexagona from the Calcaire grav-
eleux, Assise 4 Crania ignabergensis, Craie de Vend6me.
Unfortunately, the species is unrepresented in both the
BMNH and VH collections.

DISTRIBUTION. Santonian of Venddme, Loir-et-Cher,

France.

Elea labyrinthica (Michelin, 1843) Figs 57-65

1843 Eschara labyrinthica Michelin: 124, pl. 32, fig. 2.
1843 Eschara neustriaca Michelin: 124, pl. 32, fig. 3.

1853 Elea rhomboidalis d’Orbigny: 631, pl. 737, figs 21-24.
1890 Elea rhomboidalis d’Orbigny; Pergens: 399.

1899 Elea labyrinthica (Michelin); Gregory: 303.

1899 Elea rhomboidalis d’Orbigny; Gregory: 304.

MATERIAL. Type: the syntypes of this species (and of the
contemporaneous Eschara neustriaca) are not among the
Michelin types in the MNHN (see Walter 1975); Michelin
(1843) gives Cap de la Héve, Honfleur, Villers-sur-Mer and
Vaches-Noires as localities, and Craie Chloritée [= Craie
Glauconieuse, Lower Cenomanian] as the horizon. Other
material: MNHN d’Orbigny Collection 6627 [= Voigt photo-

Fig. 49 Elea hexagona d Orbigny, 1853, photograph of MNHN
d’Orbigny Collection 8192 (Voigt photocard 5791), lectotype,

- Senonian [Santonian], Vend6éme, Loir-et-Cher, France; colony is
fouled by serpulids and an oyster; x 10.

card 7475] (presumed type specimen of E. rhomboidalis),
Cenomanian, Le Havre, France; BMNH D31139, VH 10460,
Cenomanian, Cap de la Héve, Seine Maritime, France;
BMNH D58900-2, Lower Cenomanian, Craie Glauconieuse,
Cap de la Heve; BMNH D58763-4, D58890-2, Craie Glauco-
nieuse, Port d’Antifer, Seine Maritime, France; BMNH
D59156 (sample), Craie Glauconieuse, Villers-sur-Mer, Cal-
vados, France; BGS GSM 118097-8, Lower Cenomanian,
Warminster Greensand, Warminster, Wiltshire, England,
Cunnington Collection; BGS GSM Rh 4582, Cenomanian
Limestone, ?Bed Al, Hall Rocks to Beer Head, Devon,
England; BMNH D59157, Cenomanian Limestone, Bed Al
(mantelli Zone), The Pinnacles, near Beer, Devon; BMNH
D59158, Cenomanian (float), White Hart Sandpit, Wilming-
ton, Devon. VH unnumbered specimens, Lower Cenoma-
nian (carcitanensis Zone), Carriére du Billot, Notre-Dame-
de-Fresnaye, Calvados, collected by G. Breton.

DESCRIPTION. Colony bifoliate with branches of variable
morphology, strap-like, bifurcating and about 6 mm wide in
some specimens, broad and folded in others (e.g. Michelin
1843, pl. 32, fig. 2); branches about 0-9 mm deep. Zooids
arranged in regular to approximate quincunx (Figs 57, 61);
organization fixed-walled. Overgrowths extremely common,
originating through eruptive budding onto the surfaces of
erect branches; growing edges of overgrowths sometimes
closed by terminal diaphragms. Colony base extensive, giving
rise to more than one erect branch. Possible conspecific
ancestrula adjacent to a gonozooid in VH 10460 has a very
short distal tube and large protoecium about 0-22 mm wide.
Autozooids (Figs 57, 61-62) medium-sized, with frontal
walls elongate, over twice as long as wide, typically hexagonal
but occasionally diamond-shaped, pierced by circular to

Fig. 50 Elea hexagona d’Orbigny, 1853, ZMC Levinsen Collection
M39, Santonian, Vendome, France, x 28.

slightly longitudinally elongate pseudopores; zooidal bound-
aries marked by low, thread-like ridges. Apertures (Fig. 59)
small, longitudinally elongate, a little less than 1-5 x longer
than wide, attaining maximum width about mid-length,
rounded distally; apertural shelf distinct, narrow; hinge line
very slightly bowed, teeth not clearly apparent; apertural rim
raised. A large and prominent tubercle (Fig. 61) occurs
distally of the autozooidal aperture and is shaped like a
compressed barkhan dune facing towards the aperture. Oper-
cula (Fig. 58) fairly commonly found in-situ, surface convex,
bearing about 10 radial ridges peripheral to a central flatter
area; pseudopores not seen. Intramural eleozooids often
present; apertures variably D-shaped to inverted T-shaped,
located within the distal half of the aperture of the host
autozooid. Intramural autozooids possibly present but infre-
quent.

Eleozooids (Figs 57, 60, 62-64) common, scattered or in
small groups, with pseudoporous frontal walls having
approximately the same area as, though often narrower than,
those of autozooids. Apertures highly variable in length,
ranging from about 2 to 5 x longer than wide, tapering to
become very narrow at the rounded distal extremity; aper-
tural shelf wide, beginning just distally of the hinge-line;
distal tubercle absent. Opercula (Fig. 63) occasionally found
in-situ. Intramural eleozooids (Fig. 64) often present.

P. D. TAYLOR

Gonozooids known from only one specimen (VH 10460),
which has an intact and an abraded example; in the former
(Fig. 65) an initially narrow, tubular distal frontal wall
becomes densely pseudoporous before dilating into an elon-
gate ovoid shape. Ooeciopore circular or slightly longitudi-
nally elongate, about the same length as an autozooidal
aperture. Atrial ring not apparent in the abraded gonozooid,
in which the vertical walls of the more proximally overgrown
zooids protrude from the floor of the gonozooid, whereas
those of the more distal zooids do not.

MEASUREMENTS.

autozooids

(10 zooids with in-situ opercula from BGS GSM Rh 4582)
frontal length: mean = 0-55 mm; SD = 0-033 mm;
CV = 6-1; range = 0-50-0-60 mm
mean = 0-22 mm; SD = 0-023 mm;
CV = 10-2; range = 0-18-0-26 mm
mean = 0:14 mm; SD = 0-014 mm;
CV = 10-1; range = 0:12-0:17 mm
mean = 0-10 mm; SD = 0-010 mm;
CV = 10-6; range = 0-08-0-11 mm

(10 zooids from BMNH D31139)

frontal length: mean = 0-58 mm; SD = 0-043 mm;
CV = 7-4; range = 0-53-0-63 mm
mean = 0-25 mm; SD = 0-016 mm;
CV = 6:3; range = 0-23-0-27 mm
mean = 0-15 mm; SD = 0-007 mm;
CV = 4-7; range = 0:14-0:17 mm
mean = 0-11 mm; SD = 0-007 mm;
CV = 6-7; range = 0:09-0:12 mm

eleozooids (5 zooids from BGS GSM Rh 4582; 5 zooids from
BMNH D31139)
frontal length:
frontal width:
apertural length:
apertural width:

gonozooid (VH 10460)

frontal width:
apertural length:

apertural width:

frontal width:
apertural length:

apertural width:

range = 0-42-1-02 mm
range = 0-17-0-27 mm
range = 0-20-0-54 mm
range = 0-06—0-09 mm

total frontal length: 2:72 mm
distal frontal wall

length: 2:51 mm
frontal wall width: 0-98 mm
ooeciopore length: 0-11 mm
ooeciopore width: 0-11 mm

REMARKS. Although type material is lacking, the identity of
this species is not in doubt as Michelin’s enlarged figure
(1843: pl. 32, fig. 2b) shows very clearly the prominent
tubercles located distally of the autozooidal apertures. In no
other melicerititid species are the distal tubercles so well-
developed or shaped so much like barkhan dunes.

Specimens of E. labyrinthica from the type horizon — the
Craie Glauconieuse, where it is very common, exhibit vari-
able colony forms, some being adeoniform while others are
eschariform. One specimen (BMNH D58763) encrusts a
sponge.

DISTRIBUTION. Lower Cenomanian of northern France and
south-west England; known from sandy facies only.
Elea mackinneyi sp. nov. Figs 66-71

MATERIAL. Holotype: VH 10474, Lower Cenomanian,
Milheim/Ruhr, Westfalia, Germany. Paratypes: VH 10541 (4

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

Figs 51-56 Elea hexagona d’Orbigny, 1853, Santonian, Vendome, France. 51-54, ZMC Levinsen Collection M38; 51, autozooids and a
marginal eleozooid (lower left), x 33; 52, intramural operculate autozooid (left) and autozooidal aperture (right), x 135; 53, marginal
eleozooid and intramural autozooids, x 70; 54, non-marginal eleozooid, x 98. 55, 56, ZMC M39; 55, intramural autozo (left) and
normal autozooid (right), < 135; 56, autozooidal operculum, x 150.

P. D. TAYLOR

Figs 57-62 Elea labyrinthica (Michelin, 1843). 57-60, BMNH D31139, Cenomanian, Cap de la Héve, Seine Maritime, France; 5
autozooids, eleozooids and intramural eleozooids; distal tubercles abraded, x 55; 58, autozooidal operculum, x 170; 59, autozooidal
aperture, X 185; 60, eleozooid with distally broken rostrum, < 175. 61, 62, BGS GSM Rh 4582, Cenomanian Limestone, ?Bed A1, Hall

Rocks to Beer Head, Devon, England; 61, autozooids with prominent distal tubercles, x 48; 62, autozooids and eleozooids with long,
narrow rostra, X 70.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 29

Figs 63-65
Devon, England; 63, large eleozooid with slightly damaged operculum, x 145; 64, two large eleozooids housing small intramural
eleozooids, x 110. 65, VH 10460, Cenomanian, Cap de la Héve, Seine Maritime, France, gonozooid with crushed frontal wall, x 35.

specimens), same horizon and locality as holotype.

NAME. In recognition of the numerous fundamental contri-
butions made to bryozoology by Dr F.K. McKinney (Appala-
chian State University).

DESCRIPTION. Colony bifoliate with branches (Fig. 66) nar-
row, strap-like, about 1-8 mm wide between bifurcations, and
0-7 mm deep. Autozooids generally arranged in transverse
tows whose regularity is disrupted close to bifurcations;
organization fixed-walled. Overgrowths and colony base not
observed.

Autozooids (Fig. 67) medium-sized, with elongate hexago-
nal frontal walls on average slightly less than 2 x longer than
wide; pseudopores circular or longitudinally elliptical; zoo-
ecial boundaries raised. Apertures small, longitudinally elon-
gate, about 1-1 xX longer than wide, attaining maximum
width a little proximally to mid-length, appreciably narrower
at hinge line level; apertural shelf absent or at most very
slightly developed; apertural rim moderately raised; hinge
line straight or slightly bowed, typically concave in eroded

_ Zooids lacking opercula. Opercula (Fig. 69) rarely preserved

in-situ, flat or slightly depressed, bearing about 16
pseudopores arranged in a crescent parallel to the distal/
lateral margin. Terminal diaphragms not observed. Intramu-
rally budded autozooids represented by a single example with
a slightly oblique aperture (Fig. 70).

Eleozooids (Figs 67, 68) common, located along branch

_Margins, on average about 3 x longer than wide, usually

Elea labyrinthica (Michelin, 1843). 63, 64, BGS GSM Rh 4582, Cenomanian Limestone, ?Bed A1, Hall Rocks to Beer Head,

occupying two transverse rows of autozooids. Aperture long,
widest at the hinge line but broad throughout, sometimes
very slightly spatulate, well-rounded distally, on average
twice as long as wide. In-situ opercula and intramural buds
not observed.

Kenozooids (Fig. 68) occasionally present at branch mar-
gins, especially within bifurcations.

Gonozooids (Fig. 71) of unknown abundance (one fully
developed and one partially developed example present in
holotype VH 10474); distal frontal wall dilating soon after
emerging from maternal aperture, longitudinally ovoidal in
shape, about 2 x longer than wide. Ooeciopore not
observed. Atrial ring present.

MEASUREMENTS.

autozooids (10 zooids from holotype VH 10474)

frontal length: mean = 0-49 mm; SD = 0-029 mm;
CV = 6-0; range = 0-45—-0-54 mm
mean = 0:26 mm; SD = 0-018 mm;
CV = 6-7; range = 0:24-0:29 mm
mean = 0-18 mm; SD = 0-006 mm;
CV = 3-6; range = 0-17-0-18 mm
mean = 0-16 mm; SD = 0-005 mm;
CV = 2-9; range = 0-15-0-17 mm

frontal width:

apertural length:
apertural width:
eleozooids (10 zooids from holotype VH 10474)

frontal length: mean = 1:00 mm; SD = 0-184 mm;
CV = 18-5; range = 0-75-1-26 mm

Fig. 66 Elea mackinneyi sp. nov., VH 10474, holotype, Lower
Cenomanian, Milheim/Ruhr, Westfalia, Germany; branch with
broken gonozooid and fouling foraminifer (top left), x 20.

mean = 0-36 mm; SD = 0-050 mm;
CV = 13-9; range = 0-30-0-44 mm
mean = 0-49 mm; SD = 0-102 mm;
CV = 20-7; range = 0-33-0-65 mm
mean = 0-24 mm; SD = 0-034 mm;
CV = 13-8; range = 0-18-0-29 mm

frontal width:

apertural length:

apertural width:

gonozooid (from holotype VH 10474)

total frontal length: 2-34 mm
dilated frontal wall

length: 2-07 mm
frontal wall width: 1-04 mm

REMARKS. Although known from only a few specimens, this
is a very distinctive species. The broad shape of the eleozooid
rostra serve to distinguish Elea mackinneyi from sympatric
Elea elegantula sp. nov., and it differs from E. viskovae sp.
nov. from the Turonian of the Kazakhstan in having longer

rostra and in lacking an appreciable autozooidal apertural
shelf.

DISTRIBUTION. Lower Cenomanian of Germany.

P.D. TAYLOR
Figs 72-78

MATERIAL. Holotype: VH 10462, Upper Campanian, St
Severin d’Uzel, Charente Maritime, France. Paratypes: VH
10461, same horizon and locality as holotype; VH 10471,
Senonian, La gare de Soullans, Vendée, France. BMNH BZ
441-4 , Upper Campanian, Cailleau, near Talmont, Charente
Maritime, France. Other material: VH unnumbered topo-

types.

NAME. With reference to its similarity to Elea lamellosa
(d’Orbigny).

Elea pseudolamellosa sp. nov.

DESCRIPTION. Colony bifoliate, consisting of broad, folded
fronds about 0-9 mm deep. Growing edge may be occluded
by kenozooids and eleozooids. Colony base not observed.
Overgrowths may be present. Zooids arranged in approxi-
mate quincunx; organization fixed-walled.

Autozooids (Figs 72, 76) medium-sized, frontally hexago-
nal in outline, on average 1-6 X as long as wide, distal edge
prolonged by aperture; frontal wall slightly convex, with
circular pseudopores; zooecial boundary wall inconspicuous,
thin, little raised. Apertures (Fig. 74) small, longitudinally
elongate, about 1-2 x longer than wide, reaching maximum
width a little distally of the hinge line, well-rounded distally;
apertural shelf narrow; apertural rim narrow but prominent,
continuous with zooecial boundary wall; hinge line bowed.
Opercula (Fig. 73) infrequently preserved in-situ, prominent,
surface convex, seemingly with radially elongate pseudopores
widely distributed. Terminal diaphragms not observed. Intra-
mural eleozooids present, their D-shaped apertures occupy-
ing the distal part of the host aperture, which is proximally
covered by a slightly concave pseudoporous exterior wall.

Eleozooids (Figs 72, 76-77) common, often clustered,
especially near occluded growing edges; frontally variable in
size, typically only slightly smaller than autozooids; frontal
wall with circular pseudopores. Apertures (Fig. 77) small,
D-shaped, equidimensional or transversely elongate. In-situ
opercula and intramural buds not observed.

Kenozooids observed in association with eleozooids and
gonozooids (Fig. 75).

Gonozooids (Fig. 78) large, longitudinally elongate; frontal
wall with a short parallel-sided portion emerging from the
maternal zooidal aperture (Fig. 75), dilating into an ovoidal
distal portion about twice as long as wide. Ooeciopore not
observed (destroyed during abrasion of frontal wall in avail-
able specimens). Atrial ring not observed.

MEASUREMENTS.

autozooids (10 zooids from holotype VH 10462)

frontal length: mean = 0:53 mm; SD = 0-029 mm;
CV = 5-6; range = 0-48-0-57 mm
mean = 0-33 mm; SD = 0-024 mm;
CV = 7-2; range = 0:30-0:38 mm
mean = 0-18 mm; SD = 0-014 mm;
CV = 7-7; range = 0:17-0:21 mm
mean = 0-15 mm; SD = 0-007 mm;
CV = 4-7; range = 0-14-0-17 mm

from SEM micrographs of VH

frontal width:
apertural length:

apertural width:

eleozooids (estimated
10461-2)

apertural length:
apertural width:

ca 0-05—0-07 mm
ca 0-06—0-07 mm

gonozooid (one zooid from VH 10462)
frontal length: 2:52 mm

_ SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 31

oblique intramural autozooidal aperture, x 265.

2:31 mm
1:20 mm

distal frontal length:
frontal width:

REMARKS. This species is extremely similar to Elea lamellosa
d Orbigny (see p. 17). The principal point of distinction is the
D-shaped form of the eleozooid aperture, which contrasts
with the inverted T-shaped aperture characteristic of the
eleozooids in E. lamellosa. (i.e. the eleozooids of E.
pseudolamellosa are demizooids, whereas those of E. lamel-
losa are trifoliozooids; compare Figs 77 and 19) This differ-
ence appears to reflect the presence of a rostral shelf in E.
lamellosa which is absent in E. pseudolamellosa. Autozooidal
frontal walls may be a little more convex and zooecial
boundary walls rather less well-developed in E. pseudolamel-
losa than in E. lamellosa. The stratigraphical distribution of
the two species differs, E. lamellosa occurring in the
Coniacian-Santonian (and possibly the Turonian), whereas
E. pseudolamellosa is known with certainty only from the
Upper Campanian; a paratype (VH 10471) from the Vendée

Figs 67-70 Elea mackinneyi sp. nov., VH 10474, holotype, Lower Cenomanian, Milheim/Ruhr, Westfalia, Germany; 67, autozooids and
marginal eleozooid, x 55; 68, two marginal eleozooids and kenozooids at a bifurcation, x 44; 69, autozooidal operculum, x 265; 70,

assigned to E. pseudolamellosa is of unknown age within the
Senonian.

DISTRIBUTION. Upper Campanian of Charente Maritime,
and undifferentiated Senonian of the Vendée, France.

Figs 79-86

non 1846 Escharites dichotoma Reuss: 66, pl. 15, fig. 31.
1874 Echarites dichotoma (Reuss); Reuss: 135, pl. 25,
fig. 8.
1897b Elea hexagona d’Orbigny; Canu: 753.

Elea subhexagona sp. nov.

MATERIAL. Holotype: DM un-numbered, Voigt photocard
2060 (specimen figured by Reuss 1874, pl. 25, fig. 8), Upper
Turonian, ‘Oberen Planers’, Strehlen, Dresden, Germany.
Paratypes: ZMC M53-5, ‘Turonien’ [probably Cenomanian],
St Calais, Sarthe, France. Other material: MNHN Canu
Collection unnumbered.

Fig. 71 Elea mackinneyi sp. nov., VH 10474, holotype, Lower
Cenomanian, Miilheim/Ruhr, Westfalia, Germany; gonozooid
with broken frontal wall exposing atrial ring, x 48.

NAME. With reference to the resemblance to Elea hexagona.

DESCRIPTION. Colony bifoliate with branches (Fig. 79) bifur-
cating, strap-like, narrow, about 1-3—2-3 mm wide. Autozoo-
ids generally arranged in approximate quincunx; organization
fixed-walled. Colony base not observed. Overgrowth present
in ZMC M54, consisting of kenozooids spreading between
apertures of underlying autozooids, possibly free-walled (Fig.
79).

Autozooids (Figs 80, 81) medium-sized; frontal surface
usually hexagonal, elongate, about twice as long as wide;
frontal wall occupying about half of frontal surface, flat with
circular pseudopores; zocecial boundaries raised. Apertures
(Fig. 83) small, longitudinally elongate, on average
1-1-1-2 x longer than wide, attaining maximum width
between hinge line and mid-length, rounded distally; aper-
tural shelf narrow; apertural rim well-developed; hinge line
with a subdued median bar. Opercula (Fig. 82) often pre-
served in-situ, surface convex, sometimes with a flattened
median area close to the hinge line, pseudopores not visible
in studied material. Terminal diaphragms not observed.
Intramurally budded autozooids may be present, their aper-
tures typically shorter and more rounded than normal auto-
zooids.

Eleozooids (Figs 84-85) common, especially at branch
margins and in bifurcations, about 3-4 x longer than wide,
frontally slightly wider but appreciably longer than autozoo-

P! Di TAYLOR

ids. Aperture elongate, about 2-3 x longer than wide, widest
at the hinge line and tapering to a long narrow rostrum with a
rounded end. Opercula (Fig. 84) observed in-situ, surface
convex. Intramurally budded eleozooids (Fig. 81) present.

Kenozooids sometimes present in branch bifurcations.

Gonozooids (Fig. 86) with a short parallel-sided proximal
frontal wall emerging from the maternal zooid aperture,
becoming longitudinally ovoidal, distal dilated frontal wall
approximately 1-5 x longer than wide. Ooeciopore removed
by abrasion; atrial ring present.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from the holo-
type)

frontal length: mean = 0:53 mm; SD = 0-040 mm;
CV = 7-6; range = 0-48-0-60 mm
mean = 0:27 mm; SD = 0-019 mm;
CV = 7-1; range = 0-26-0-30 mm
mean = 0-19 mm; SD = 0-007 mm;
CV = 4-1; range = 0-18-0-20 mm
mean = 0-17 mm; SD = 0-007 mm;
CV = 4:3; range = 0-15—0-18 mm

frontal width:
apertural length:

apertural width:

eleozooids (5 zooids from the holotype)

frontal length: range = 0-74-0-81 mm
frontal width: range = 0:23-0:29 mm
apertural length: range = 0-36-0-42 mm
apertural width: range = 0-15—0-20 mm

gonozooids (2 zooids estimated from SEM micrographs of
ZMC MS3 and M55)
total frontal length:
distal frontal wall
length*: ca 1-24-1-44 mm

frontal width: ca 0:85-0:96 mm

(* i.e. that part of the frontal wall distal to the narrow origin)

ca 1-61-1-88 mm

REMARKS. This new species was identified by Reuss (1874)
as Escharites dichotoma Reuss, 1846. Unfortunately, the type
specimen of E. dichotoma, from the Exogyrensandstein of
Bohemia, was destroyed in 1956 (E. Voigt pers. comm.
December 1986). However, it is clear from the figure of
Reuss (1846), which depicts a vinculariiform colony resem-
bling Filicea, that it is not the same species as E. dichotoma —
sensu Reuss, 1874.

Canu (1897b) identified specimens here regarded as‘Elea |
subhexagona as E. hexagona d’Orbigny, a Santonian species
which is very similar to E. subhexagona but which can be
distinguished by the deep and prominent tubercle situated |
distally to the autozooidal aperture. Canu apparently realized |
later that this identification was mistaken because material of
his in the MNHN and in the ZMC (donated to G.M.R.
Levinsen) is labelled with the MS name ‘Elea fissurata’. The |
exact stratigraphical horizon of these specimens from St |
Calais (Sarthe) is questionable; Canu originally ascribed his |
St Calais material to the Cenomanian but subsequently |
referred it to the Turonian (Canu and Bassler, 1922). It seems |
possible that the specimens are from the late Cenomanian |
plenus Zone, although E. subhexagona is not represented |
among material of this age from St Calais in the VH and
BMNH collections.

DISTRIBUTION. Upper Cenomanian/Lower Turonian of St }
Calais, Sarthe, France, and Upper Turonian of Dresden,
Germany. |

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 33

Figs 72-77 Elea pseudolamellosa sp. nov. 72-74, VH 10461, Upper Campanian, St Severin d’Uzel, Charente Maritime, France; 72,
autozooids and eleozooids (top left), x 47; 73, autozooidal operculum, x 250; 74, autozooidal aperture, x 250. 75, VH 10462, holotype,
Upper Campanian, St Severin d’Uzel, gonozooid origin with kenozooid above and to the left of the maternal zooid, x 65; 76, 77, VH
10471, Senonian, Le gare de Soullans, Vendée, France; 76, eleozooids and autozooids, x 65; 77, apertures of two eleozooids, x 250.

Fig. 78 Elea pseudolamellosa sp. nov., VH 10462, holotype, Upper
Campanian, St Severin d’Uzel, Charente Maritime, France;
gonozooid with broken roof, x 42.

Elea triangularis (Michelin, 1841) Figs 87-94

1841 Eschara triangularis Michelin: 5, pl. 1, fig. 6.

1850 Eschara triangularis Michelin; d’Orbigny: 140.

1851 Eschara triangularis Michelin; d’Orbigny, pl. 602, figs
4 and S.

1853 Elea triangularis (Michelin); d’Orbigny: 630, pl. 737,
figs 17-20.

1890 Elea triangularis (Michelin); Pergens: 397.

1899 Elea triangularis (Michelin); Gregory: 305.

1975 Elea triangularis (Michelin); Walter: 314, pl. 29, figs 1,
>:

MATERIAL. Holotype: _MNHN _ Michelin Collection
un-numbered (Fig. 87), Albian, Grandpré, Ardennes,
France. Other material: MNHN d’Orbigny Colln 6020 (5
pieces mounted on a tablet with the mark of a sixth, lost
piece), same horizon and locality as holotype; BMNH 60541
(large colony and several fragments), ‘Gault Chloritic’ (prob-
ably Lower Albian mammillatum Zone according to H.G.
Owen pers. comm.), Folkestone, Kent, UK; BMNH D52080
(SEM stub), Lower Albian (tardefurcata or mammillatum
Zone, see Owen, 1972; Rawson et al., 1978), Shenley Lime-
stone, Leighton Buzzard, Bedfordshire, UK, R.J. Hogg
Collection.

DESCRIPTION. Colony bifoliate (Fig. 87) with narrow, folded
branches which bifurcate occasionally; branches generally 6

P. D. TAYLOR

Fig. 79 Elea subhexagona sp. nov., ZMC Levinsen Collection
M54, ‘Turonien’ [probably Cenomanian], St Calais, Sarthe,
France; branch with autozooids, eleozooids and distally, an
overgrowth of kenozooids occupying the areas between zooidal
apertures; x 39.

mm wide and 1 mm deep. Zooids arranged in approximate
quincunx; organization fixed-walled. Colony base hollow (in
holotype). Growing edge often arcuate. Overgrowths some-
times developed on erect branches. Zooidal buds may be
occluded by terminal diaphragms located well proximal to the
interzooidal wall ends.

Autozooids (Fig. 89) medium-sized, generally rhombic
with relatively short, convex frontal walls pierced by large,
circular, slightly countersunk pseudopores; zooidal bound-
aries raised. Apertures (Fig. 90) very large, elongate, almost
1-5 x longer than wide, attaining maximum width a little
distal to the hinge line, well-rounded distally; apertural shelf
present only in the mid-part of the aperture, indenting the
margins of the aperture and producing the bell-shape typical
of the species; hinge line with a pair of prominent hinge teeth.
Apertural rim thick, continuous with variably extensive but
low tubercles at the lateral corners of the frontal wall.
Opercula (Fig. 88) occasionally preserved in-situ, proximal
edge a little concave, surface slightly convex, fissured and
with about 25 elongate pseudopores arranged in an irregular
crescent; internally (Fig. 92), prominent, inward-sloping
sclerites form a continuous arch immediately outward of the
pseudopores. Intramurally budded autozooids may be
present. A tubular structure apparently composed of

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

WwW
n

Figs 80-83 Elea subhexagona sp. nov. 80, DM un-numbered, Voigt photocard 2060, holotype, Upper Turonian, “Oberen Planers’, Strehlen,
Dresden, Germany, autozooids, mostly with opercula, and eleozooids, x 45. 81-83, ZMC Levinsen Collection M54, ‘Turonien’ [probably
Cenomanian], St Calais, Sarthe, France; 81, autozooids and a primary eleozooid containing an intramural eleozooid, x 70; 82, autozooidal
operculum, X 210; 83, autozooidal aperture, x 210.

pseudoporous exterior wall, which can be seen within the
aperture of one zooid (Fig. 91), is interpreted as a bioclaus-
tration (i.e. an embedment structure formed by the bryo-
zoan; cf. similar structures in Cretaceous onychocellids
attributed to the presence of folliculinid protists by Ernst,
1985). Interzooidal walls with conspicuous pores. Mural
spines present in great abundance in the proximal parts of
zooids, long, and either simple or branched (Fig. 93).

Eleozooids unknown, presumed absent.

Kenozooids sometimes present.

Gonozooid known from only one partly developed
example (Fig. 94) showing the tubular, densely pseudoporous
proximal part of the frontal wall emerging from the aperture
of the maternal zooid.

MEASUREMENTS.

autozooids (10 zooids from BMNH 60531)

frontal length: mean = 0-55 mm; SD = 0-036 mm;
CV = 6-6; range = 0-50-0-60 mm

frontal width: - mean = 0-35 mm; SD = 0-027 mm;

CV = 7:8; range = 0-32-0-39 mm
mean = 0:29 mm; SD = 0-015 mm;
CV = 5-2; range = 0:27-0:32 mm
mean = 0-20 mm; SD = 0-011 mm;
CV = 5:2; range = 0-20-0-23 mm

apertural length:

apertural width:

REMARKS. The large size and characteristically bell-shaped
autozooidal apertures immediately distinguish this uncom-
mon species from others in the genus. E. triangularis is the
only bifoliate melicerititid described from the Lower Creta-
ceous. It is also currently unique in being the only melicer-
ititid known to possess mural spines (= zooecial or
intrazooecial spines) (Fig. 93). Similar structures are, how-
ever, common and widely distributed taxonomically among
non-melicerititid cyclostomes (see Farmer, 1979) and further
investigations of the internal morphology of melicerititids will
probably reveal more examples.

DISTRIBUTION. Albian (?Lower Albian only) of France and
England.

36 P. D. TAYLOR

Figs 84-86 Elea subhexagona sp. nov. 84, DM un-numbered, Voigt photocard 2060, holotype, Upper Turonian, “Oberen Planers’, Strehlen,
Dresden, Germany, eleozooid with operculum, x 145. 85, 86, ZMC Levinsen Collection M53, ‘Turonien’ [probably Cenomanian], St
Calais, Sarthe, France; 85, eleozooid, x 120; 86, damaged gonozooid, x 53.

Fig. 88 Elea triangularis (Michelin, 1841), BMNH D52080, Lower

87 Albian (tardefurcata or mammillatum Zone), Shenley Limestone,
Fig. 87 Elea triangularis (Michelin, 1841), photograph of the Leighton Buzzard, Bedfordshire, UK; secondary electron image
holotype colony, MNHN Michelin Collection un-numbered, of coated specimen showing autozooidal operculum in slightly

Albian, Grandpré, Ardennes, France; x 2-5. oblique view, x 117.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

| Figs 89-94 Elea triangularis (Michelin, 1841), BMNH 60531, ‘Gault Chloritic’ [probably Lower Albian mammillatum Zone], Folkestone,
| Kent, UK; 89, group of autozooids, including one with an intramural autozooid (upper centre), < 62; 90, autozooidal aperture, x 125; 91,
autozooidal aperture containing enigmatic tubular structure, x 120; 92, inner side of an autozooidal frontal wall and operculum showing
sclerites and pseudopores, x 118; 93, fractured specimen exposing mural spines within the proximal chamber of an autozooid, x 355; 94,
proximal part of a broken gonozooid, x 37.

38
Figs 95-99

MATERIAL. Holotype: VH 10450, Turonian (‘C22’ division),
Kyzylsaj, ca 180 km east of Fort Shevchenko, Kazakhstan,
Najdin Collection. Paratypes VH 10542 (6 specimens),
details as for holotype.

Elea viskovae sp. nov.

NAME. In honour of the Russian bryozoologist L.A.
Viskova.

DESCRIPTION. Colony bifoliate with branches (Fig. 95) nar-
row, strap-like, bifurcating, about 1-3-2-0 mm wide and
0-5—0-6 mm deep. Autozooidal arrangement variable, both
autozooids and eleozooids divergent close to branch margins;
organization fixed-walled. Overgrowths common. Colony
base unknown. One of the paratypes grows against and
partially envelops a ?serpulid tube.

Autozooids (Fig. 96) large; frontal walls irregularly hex-
agonal, distally rounded, elongate, 2-5-3 = longer than wide,
with circular pseudopores; zooecial boundaries raised. Aper-
tures (Fig. 99) of moderate size, longitudinally elongate, on
average 1-1 x longer than wide, attaining maximum width
just proximal to mid-length, appreciably narrower at hinge
line level, well-rounded distally; apertural shelf broad,
0:02-0:03 mm wide, causing the aperture to have the shape of
a high isosceles triangle with rounded corners; apertural rim

Fig.95 Elea viskovae sp. nov., VH 10450, holotype, Turonian
((C22’ division), Kyzylsaj, ca 180 km east of Fort Shevchenko,
Kazakhstan, Najdin Collection; branch with autozooids and
marginal eleozooids, x 37.

P: DI TAYEOR

moderately prominent; hinge line bowed. Opercula (Fig. 97)
often preserved in-situ, slightly convex, with about 16
pseudopores arranged in a cresent parallel to the distal/lateral
margin. Terminal diaphragms (Fig. 97) positioned proximally
to the apertural shelf, ?non-pseudoporous. Intramural buds
not observed with certainty.

Eleozooids (Figs 95, 96) common, located close to branch
margins, about 2—2-5 longer than wide, their frontal walls a
little longer and significantly wider than those of the autozoo-
ids. Aperture elongate, attaining maximum width slightly
distally to the hinge-line, well-rounded distally; apertural
shelf not clearly developed but rostral shelf present. Opercula
known from only one partial example in-situ, surface strongly
convex at preserved edges. Intramural buds not observed.

Kenozooids commonly present at branch margins.

Gonozooids not observed.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from holotype VH
10450)
frontal length: mean = 0-64 mm; SD = 0-070 mm;
CV = 10-9; range = 0:54-0:77 mm
mean = 0-24 mm; SD = 0-024 mm;
CV = 9-6; range = 0-21-0-29 mm
mean = 0-19 mm; SD = 0-009 mm;
CV = 4-4; range = 0-18-0-21 mm
mean = 0-17 mm; SD = 0-010 mm;
CV = 6-0; range = 0-17-0-20 mm

eleozooids (5 zooids from holotype VH 10450)
frontal length: range = 0-63—0-80 mm
frontal width: range = 0-32-0-35 mm
apertural length: range = 0-27-0-30 mm
apertural width: range = 0-21-0-24 mm

frontal width:
apertural length:

apertural width:

REMARKS. The marginal eleozooids of Elea viskovae, which
resemble slightly-enlarged autozooids, distinguish this new
species from others within the genus Elea. They have less
well-developed rostra than those of E. mackinneyi, a species
with smaller autozooids. Eleozooid morphology recalls that
present in such species as Meliceritites semiluna and
Biforicula filicosa.

Many of the autozooidal opercula (and some terminal
diaphragms) in the holotype are pierced by small circular
borings (Figs 96-97) like those described previously in other
melicerititids (Taylor, 1982).

DISTRIBUTION. Turonian of Kazakhstan.

Figs 100-109

MATERIAL. Holotype: VH 10538, top Lower Cenomanian
(orbignyi Zone) — basal Middle Cenomanian (costatus Zone),
Saint-Germain-la-Campagne, descente sur Orbiquet, Eure,
France, collected by G. Breton 1981. Paratypes: VH 10439,
10539-40, same horizon and locality as holotype. BMNH
D55027-8, Cenomanian [?Lower Cenomanian, Craie Glau-
conieuse], Bruneval, Seine-Maritime, France, S. Whiteley
Collection. Other material: VH, numerous un-numbered
topotypes.

Elea whiteleyi sp. nov.

NAME. In honour of the late S. Whiteley, an enthusiastic
amateur geologist who collected paratypes of this species.

DESCRIPTION. Colony bifoliate with bifurcating, narrow
strap-like branches about 1-1—2-3 mm wide and 0-6 mm deep,
elliptical in cross-section. Autozooids arranged more or less

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 39

in quincunx or in ill-defined transverse rows (Fig. 100),
separated from adjacent autozooids by demizooids; organiza-
tion fixed-walled. Overgrowths moderately common, at least
some formed by eruptive budding onto surfaces of erect
branches, sometimes consisting predominantly of demizooids
and kenozooids . Colony base (Figs 105-107) extensive,
giving rise to several erect branches whose median budding
laminae are parallel to local orientation of basal zooids;
demizooids of base have higher, more pointed apertures than
those of erect branches, and many possess non-inclined
apertures orientated parallel to adjacent autozooids.
Autozooids (Figs 102, 104) small, with longitudinally ellip-
tical pseudoporous frontal walls, about twice as long as wide,
depressed with respect to the surrounding demizooids. Aper-
tures small, longitudinally elongate, about 1-1 x longer than
wide, attaining maximum width at a level between the hinge
line and mid-length; apertural shelf slight; apertural rim
prominent, sometimes prolonged at the mid-point of the
aperture; hinge line with a curved median bar. Opercula (Fig.
101) very often preserved in-situ, surface moderately convex,
a flattened median proximal area often evident, bearing an
estimated 14~18 slit-shaped pseudopores in a crescent close to

—
Re

Ce gs
rae

Figs 96-99 Elea viskovae sp. nov., VH 10450, holotype, Turonian (“C22’ division), Kyzylsaj, ca 180 km east of Fort Shevchenko,
Kazakhstan, Najdin Collection; 96, autozooids with opercula, some bored, and three marginal eleozooids, x 62; 97, bored autozooidal
operculum (upper left) and terminal diaphragm (lower right), x 160; 98, autozooidal operculum, x 250; 99, autozooidal aperture, 250.

the lateral/distal edge. Intramural buds and terminal dia-
phragms not observed.

Rostrozooids (Figs 105, 108) moderately common, usu-
ally occurring singly close to the branch mid-line just
before a bifurcation, but sometimes located close to branch
margins, elongate, about 3-5 x longer than wide. Aper-
ture long, attaining maximum width at the hinge line, very
slightly spatulate, rounded distally, about 3-4 x longer
than wide. In-situ opercula not seen. Rostrozooids and/or
demizooids may be budded intramurally within rostrozoo-
ids (Fig. 108).

Demizooids (Fig. 103) numerous, normally about 2 per
autozooid, often paired disto-laterally on either side of an
autozooidal aperture; frontal walls usually long and narrow,
forming raised areas between frontal walls of laterally adja-
cent autozooids, their boundaries, however, poorly-defined;
apertures variously orientated, many in oblique or reverse
orientation with respect to colony growth direction, typically
in a plane inclined about 60° to the colony surface, small,
semicircular to longitudinally elongate in shape, rounded or
slightly arched distally. Opercula often preserved in-situ;
terminal diaphragms observed.

PDF TAWIEOR

Figs 100-103 Elea whiteleyi sp. nov., top Lower Cenomanian (orbignyi Zone) — basal Middle Cenomanian (costatus Zone),
Saint-Germain-la-Campagne, descente sur Orbiquet, Eure, France. 100-101, VH 10539; 100, branch surface, x 53; 101, autozooidal
operculum, x 330. 102, 103, VH 10538, holotype; 102, autozooids, kenozooids, and demizooids with obliquely-directed, inclined
apertures, X 95; 103, operculate autozooid and two demizooids, x 220.

Kenozooids (Figs 102, 105) observed commonly in colony
base, overgrowths, and at the margins of erect branches.

Gonozooids (Fig. 109) fairly common. Distal frontal wall
emerging from maternal aperture initially parallel-sided
before becoming bulbous and longitudinally ovoidal in shape.
Ooeciopore located beyond inflated part of frontal wall,
transversely elliptical, about 1-5 x wider than long. Atrial
ring not observed.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from holotype VH
10538)

frontal length: mean = 0-33 mm; SD = 0-015 mm;
CV = 4-5; range = 0:30-0:35 mm
mean = 0:17 mm; SD = 0-008 mm;
CV = 4-5; range = 0-17-0-18 mm
mean = 0-15 mm; SD = 0-005 mm;
CV = 3-1; range = 0:15-0:17 mm
mean = 0:13 mm; SD = 0-006 mm;
CV = 4-8; range = 0-12-0-:14 mm

frontal width:
apertural length:

apertural width:

rostrozooids (5 zooids from 5 colonies)

frontal wall length: range = 0-63-0-92 mm
frontal wall width: range = 0-20-0-21 mm
apertural length: range = 0-33-0-57 mm
apertural width: range = 0-11-0-15 mm

demizooids

apertural width: ca 0-06-0-09 mm

gonozooids (one zooid from holotype VH 10538)
total frontal wall

length: 2-36 mm
dilated frontal wall

length*: 1-13 mm
ooeciopore length: 0-08 mm
ooeciopore width: 0-12 mm

(* i.e. that part of the frontal wall distal to the narrow origin)

REMARKS. The presence of numerous small eleozooids
enables this new species to be readily distinguished from all
other species of Elea, including E. elegantula which it most

resembles in the morphology of the other polymorphs. These
' eleozooids and their distribution surrounding the autozooids
invite comparison between E. whiteleyi and the Upper Cen-
/omanian bifoliate melicerititid Biforicula multicincta, the
‘oldest known species of Biforicula (see Voigt, 1989) The
principal qualitative difference between E. whiteleyi and B.
_multicincta is the existence of cancelli in the latter. It there-
' fore seems possible that E. whiteleyi belongs in the stem-
| group of Biforicula.

DISTRIBUTION. Lower-Middle Cenomanian of Normandy,
France.

Genus SEMIELEA d’Orbigny, 1853

TYPE SPECIES. Entalophora vieilbanci d’Orbigny, 1850, des-
ignated by Bassler (1935: p. 198) as ‘S. vielbanci D’Orbigny,
1853’ [sic].

| SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 4]

Figs 104-107 Elea whiteleyi sp. nov. 104, BMNH D55027 Cenomanian [?Lower Cenomanian, Craie Glauconieuse], Bruneval,
Seine-Maritime, France, operculate autozooids (demizooids obscured by diagenetic cement), x 98. 105-107, VH 10439, top Lower
Cenomanian (orbignyi Zone) — basal Middle Cenomanian (costatus Zone), Saint-Germain-la-Campagne, descente sur Orbiquet, Eure,
France; encrusting colony base; 105, irregular arrangement of autozooids, kenozooids, small eleozooids and a large eleozooid, x 40; 106,
large eleozooid (rostrozooid), x 100; 107, operculate autozooid and small eleozooid, x 135.

OTHER SPECIES. Semielea dichotoma d@’Orbigny, 1853.

REVISED DIAGNOSIS. Eleid with tubular colony-form; axial
lumen of subequal diameter lined by an inferred exterior
wall; zooids in primary layer orientated parallel to tube
length; autozooids fixed-walled; cancelli lacking.

REMARKS. D’Orbigny (1853) referred three species to Semie-
lea. Only the first two are here retained in the genus; the
third, Semielea plana d’Orbigny, 1853 is regarded as the base
of Elea lamellosa (d’ Orbigny, 1850) (see p. 19).

Diagnostic of Semielea is the possession of branches with
an axial lumen, the bounding wall of which forms a lamina for
zooidal budding. The axial lumen is intermediate in size
between the narrow axial canals present in some species of
Meliceritites (e.g. M. semiclausa (Michelin)) and the wide
tubular interiors of certain specimens of Reptomultelea with
cavariiform colonies. The growth of the axial lumen in
Semielea also seems distinct. Although no specimens have

P.D. TAYLOR

Fig. 108 Elea whiteleyi sp. nov., VH 10539, top Lower
Cenomanian (orbignyi Zone) — basal Middle Cenomanian
(costatus Zone), Saint-Germain-la-Campagne, descente sur
Orbiquet, Eure, France; primary rostrozooid containing
intramural rostrozooid in turn containing intramural
demizooid, x 120.

been studied with their microstructure sufficiently well-
preserved to prove that the wall which lines the lumen is an
exterior wall rather than an interior wall, the transverse
folding and wrinkling of this wall (Fig. 119) strongly suggests
that it is an exterior wall. Rare examples of colony bases
viewed from the underside (Fig. 122) show that the lumen of
each branch opens at the colony base and that the lumen wall
is smoothly continuous with the basal lamina. As the basal
lamina is an undoubted exterior wall, this observation is
further evidence that the lumen walls are exterior. In S.
vieilbanci occasional transverse platforms partition the axial
lumen and seem to have been formed by progressive constric-
tion of the lumen. The presence of these platforms, which
would have periodically sealed the entrance to the lumen at
branch growth tips, may explain why fouling of the apparent
exterior walls lining the lumen has not been observed (cf.
exterior walls lining the tubes of cavariiform Reptomultelea
colonies). The platforms also imply that branches could not
have grown around another erect organism (cf. some species
of Reptomultelea), but instead grew freely into space.

The phylogenetic relationship is unclear between Semielea
and species of Meliceritites on the one hand and of Reptomul-
telea on the other. Assuming Semielea to be a relatively
advanced genus, it could have derived from either of the two
latter genera. Derivation from Meliceritites would necessitate

Fig. 109 Elea whiteleyi sp. nov., VH 10538, holotype, top Lower
Cenomanian (orbignyi Zone) — basal Middle Cenomanian
(costatus Zone), Saint-Germain-la-Campagne, descente sur
Orbiquet, Eure, France; gonozooid, x 50.

the incorporation of tubular prolongations of the basal lamina
into the centre of the erect cylindrical branches, together with
loss of the normal endozonal budding pattern. Derivation
from Reptomultelea would require formation of tubular pro-
longations of the basal lamina to give erect cylindrical
branches. On the grounds of parsimony, the latter alternative
seems the more likely, but future phylogenetic analysis,
including zooid-level characters, might provide better evi-
dence of affinity.

DISTRIBUTION. Turonian (?Upper) — Lower Santonian,
northern France.

Semielea vieilbanci (d’Orbigny, 1850)

1850 Entalophora vieilbanci d’Orbigny: 200.

1851 Diastopora vieilbanci (d’ Orbigny); d’Orbigny, pl. 637,
figs 7-8.

1851 Diastopora arborescens d’Orbigny: pl. 638, figs 1-5.

1853 Semielea vieilbanci (d’Orbigny); d’Orbigny: 636, pl.
738, figs 5—9.

1853 Semimultelea arborescens (d’Orbigny); d°Orbigny:
652, pl. 741, fig. 5.

1890 Semielea vieilbanci (d’Orbigny); Pergens: 392.

1893 Semielea vieilbanci (d’Orbigny); Pergens: 211.

21897a Semielea vieilbanci (d’Orbigny); Canu: 155.

21897b Semielea vieilbanci (d’Orbigny); Canu: 750.

1899 Elea vieilbanci (d’Orbigny); Gregory: 300, fig. 33.

Figs 110-122

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 43

Fig. 110 Semielea vieilbanci (d’Orbigny, 1850), photograph of
MNHN d’Orbigny Collection 6965 (part), Turonian,
France; x 8-S.

1912 Meliceritites vieilbanci (d’Orbigny); Levinsen: 47, pl.
1, figs 15 and 16.

1953 Semielea vielbanci [sic] (d’Orbigny); Bassler: G77, fig.
40, 3a, b.

1981 Semielea vieilbanci (d’Orbigny); Voigt: 453, fig. 5a.

MATERIAL. Type: there is no material of this species from
the type locality (Turonian of Tourtenay; see d’Orbigny 1850,
p. 200) in the d’Orbigny Collection, MNHN, although over a
hundred specimens from the Turonian of several other locali-
ties are registered under number 6965 (e.g. Fig. 110). In view
of the absence of topotype specimens, together with the
unequivocal identity of the species, no neotype designation is
made.

Other material: MNHN d’Orbigny Colln 6965, over 100
pieces from the Turonian of various localities in France,
including Troot, St Maure, Tours, Angouleme, Villavard.
MNHN d’Orbigny Colln 8200 (Figs 121, 122), Senonian,
?Villedieu, ?Vendéme, syntypes of Diastopora arborescens
d’Orbigny, 1851, several specimens including Voigt Photo-
card 8312 and an un-numbered specimen corresponding to
that illustrated by d’Orbigny (1851: pl. 638, fig. 2). ZMC
Levinsen Colln unnumbered, Bruillé Poncé and Villedieu,
France. VH unnumbered: U. Turonian, Souge; Turonian,
Les Ulmes, Eure; U. Turonian, Bois de Gareau, nr Ecom-
moi, Sarthe; U. Turonian, Angoumien, La Charte sur la
Loire; U. Turonian, Sougé, Loir-et-Cher; Turonian, Luynes,
Indre et Loire; Turonian, Les Ulmes, Maine; U. Turonian,

_Troot, Loir-et-Cher; U. Turonian, Les Roches, Loir-et-Cher;
-Turonian, Les Ulmes, Maine; U. Turonian/Coniacian, La

Barre, nr St Maure, Indre et Loire; Coniacian, St Paterne,
Indre et Loire; Coniacian, Tours, Indre et Loire; Coniacian,
Fécamp, Seine Maritime. BMNH D3747 (4), ‘Senonian’,
Lavardin, Gamble Colln; D8959-60, ‘Senonian’, Lavardin;
D4807, Craie de Villedieu [Coniacian or Santonian] , S. of les
Roches; D4672, Craie de Villedieu, Luynes; D4679, D4800,
D4806, D4860 (4), [Turonian], Montloire; D4675, D4676 (5),
D4677, D4678 (4), D4683 (6), D4684 (3), D4715, D4799,
D4811 (2), D4839, D4840 (3), D4861, D4862 (5), D4864 (2),
D4865, D4868, D13573, Craie Marneuse, Villardin; D4937,
[?Turonian], “Nodula Chalk’, Villardin Castle, purchd F.H.
Butler; D4674 (figd Gregory, 1899: fig. 33), D4685, Craie
Marneuse, Chinon; D33939-40, Turonian, St Calais, Sarthe,
E. Darteville Colln; D36151—5, ‘Senonian’, Lavardin, Loir-
et-Cher, Pergens Colln; D36156, Turonian, Lavardin, Loir-
et-Cher, Pergens Colln; D53444, Turonian, Les Ulmes,
Maine, Voigt Colln; D59368, Santonian, Crate de Villedieu,
Bouchardiére Mbr, Bed 20 of Jarvis et al. (1982), La Boucha-
rdiére, Villedieu, MLoir-et-Cher; D59369 (3), Craie de
Villedieu [Coniacian or Santonian], Luynes; D59370-2,
D59373 (3), D59374 (6), D59375 (sample), Turonian, Lavar-
din, Loir-et-Cher, F. Canu Colln; D59376 (6), Turonian,
Chatellerault, Vienne, France; BZ 729 (sample), Turonian,
St Maure, Indre-et-Loire, France, presd E. Voigt, November
1991.

DESCRIPTION. Colony tubular (cavariiform) with branches
(Fig. 110) bifurcating, normally subcircular in cross-section
but becoming elliptical immediately prior to bifurcation,
varying from about 1-0 to 4-4 mm in diameter, typically
3-5-4-5 mm. Axial lumen lined by transversely wrinkled,
inferred exterior wall (Fig. 119) and generally 0-6—-1-4 mm in
minimum diameter, becoming elliptical in cross section prior
to dividing at branch bifurcations. Single layer thickness
about 0-3-0-5 mm. Budding apparently occurs only on the
lumen-lining wall. Overgrowths occasionally present, origi-
nating through intrazooecial fission, either disc-shaped and
multidirectional or fan-shaped and unidirectional (Fig. 118);
pseudoancestrula an autozooid, aperture about 0-09 mm long
by 0-08 mm wide, initiating a secondary zone of astogenetic
change of increasing zooid size. Zooid apertures arranged in
rough quincunx or in rows transverse to growth direction.
Organization fixed-walled.

Autozooids (Figs 111, 115) of moderate size, frontally
elongate, subhexagonal or subrhomboidal in outline, on
average 1-7—1-8 < longer than wide, well-rounded distally;
boundary wall salient, forming a widened wall with contigu-
ous apertural rim at distal end of zooid; frontal wall flat
proximally, sometimes slightly convex distally, pseudopores
subcircular. Apertures (Fig. 113) small, occupying less than a
third of the frontal area, approximately equidimensional or a
little wider than long, attaining maximum width about mid-
length, well-rounded distally; apertural rim salient; apertural
shelf of moderate width, tapering proximally; hinge line
poorly preserved in all scanned specimens, bowed. Opercula
(Fig. 112) often preserved in-situ, convex, prominent;
pseudopores radially elongate, numbering about 16, arranged
in a crescent. Terminal diaphragms (Fig. 117) common,
located just proximally to the apertural shelf, with a central
depression and pore. Intramurally budded autozooids not
observed. Intramurally budded eleozooids (Fig. 111) very
common; aperture (Fig. 114) trifoliate, prominent.

Kenozooids (Fig. 116) uncommon, developed close to
branch bifurcations.

P. D. TAYLOR

Figs 111-114 Semielea vieilbanci (d’Orbigny, 1850), BMNH D59370, Turonian, Lavardin, Loir-et-Cher, France; 111, group of autozooids,
some containing intramural eleozooids, X 75; 112, autozooidal operculum, x 330; 113, autozooidal aperture, <x 330; 114, intramural
eleozooidal aperture, x 365.

Eleozooids (Fig. 116) rare (excluding intramurally budded
examples); frontally about the same length but narrower than
the autozooids, pointed distally; aperture trifoliate; opercula
not observed.

Gonozooids (Fig. 120) rare; frontally elongate, about twice
as long as wide, the inflated frontal wall emerging from the
maternal aperture initially parallel-sided, becoming longitudi-
nally ovoidal distally: Ooeciopore and atrial ring not seen in
poorly preserved specimens available for study.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from BMNH
D59370)
frontal length: mean = 0:47 mm; SD = 0-030 mm;
CV = 6-4; range = 0-42-0-53 mm
mean = 0-27 mm; SD = 0-019 mm;
CV = 6:9; range = 0-26-0-30 mm
mean = 0:13 mm; SD = 0-007 mm;
CV = 5-5; range = 0:12-0:14 mm
mean = 0:14 mm; SD = 0-009 mm;
CV = 6:3; range = 0:12-0:15 mm

frontal width:
apertural length:

apertural width:

eleozooids (1 zooid from BMNH D36151)

frontal length: 0-48 mm
frontal width: 0-20 mm
apertural length: 0-09 mm
apertural width: 0-04 mm

gonozooids (2 zooids from BMNH D4674)

total frontal length: 1-86-1-91 mm
dilated frontal wall

length: 1:56-1:65 mm
frontal width: 0:90-0:98 mm

REMARKS. This is a very common species. Most specimens
come from the Upper Turonian of the Loire region of France,
but the species ranges upwards into the Coniacian and
evidently even into the basal Santonian. The wide axial
lumen immediately distinguishes S. vieilbanci from similarly
dendroid species of Meliceritites which, at most, only have
narrow axial canals a little wider than the zooecial chambers.
The small size of the apertures, particularly in comparison
with frontal wall dimensions, distinguishes S. vieilbanci from

;
;
’
)

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 45

Figs 115-118 Semielea vieilbanci (d’Orbigny, 1850). 115-117, BMNH D36151, ‘Senonian’, Lavardin, Loir-et-Cher, France; 115, autozooids
arranged in transverse rows, X 60; 116, autozooids, kenozooids and an eleozooid (centre) close to a branch bifurcation, x 100; 117,
autozooidal aperture with terminal diaphragm, x 340. 118, BMNH D59371, Turonian, Lavardin, overgrowth origin, x 135.

S. dichotoma, and both species of Semielea differ from
cavariiform colonies of Reptomultelea spp. in the consistently
parallel orientation of the long axes of the zooids and the
tubular branches.

DISTRIBUTION. Turonian (?Upper) — Lower Santonian,
northern France.

Semielea dichotoma (d’Orbigny, 1851)

1851
1853
1890
1912

Figs 123, 124

Diastopora dichotoma d’Orbigny: pl. 638, figs 6-8.
Semielea dichotoma (d’Orbigny); d’Orbigny: 637.
Semielea dichotoma (d’Orbigny); Pergens: 393.
Meliceritites dichotoma (?) (d’Orbigny); Levinsen: 38,
pl. 5, figs 1 and 2.

MATERIAL. Lectotype: MNHN d’Orbigny Collection 8194
(Figs 123-124), specimen labelled as ‘type’ by E. Voigt, Voigt
Photocard No. 8321, Senonian [?Coniacian], Tours, Indre et

Loire, France. This specimen, a fragment 21 mm long which
resembles but does not exactly match plate 638, figure 6 of
d’Orbigny (1851), is one of several specimens of various
species contained in six tubes and registered as 8194. No
other material is available for study. E. Voigt (in litt. April
1984) states that ‘Although I have got much material from
Tours, I have never found this species in the French Turo-
nian’.

DESCRIPTION. Colony tubular (cavariiform) with bifurcating
branches (Fig. 123) about 2-3 mm in diameter. Overgrowths
not observed. Organization fixed-walled. Zooid apertures
variously arranged, often irregular.

Autozooids (Fig. 124) with short frontal walls. Apertures
large, frontally elongate or equidimensional, closely-spaced,
well-rounded distally; apertural rims thick and protruberant,
especially distally; apertural shelf broad. Opercula often
preserved in-situ, strongly convex and prominent. Terminal

Fig. 119 Semielea vieilbanci (d’Orbigny, 1850), BMNH D59372,
Turonian, Lavardin, Loir-et-Cher, France; longitudinally
fractured branch showing axial lumen with wrinkled lining
wall, x 21.

diaphragms with a transverse depression present in many
zooids. Intramurally budded eleozooids present; aperture
triangular in shape, longer than wide.

Kenozooids not observed.

Eleozooids (except intramural buds) of uncertain presence.

Gonozooids not observed.

MEASUREMENTS (approximate determinations taken from
the lectotype MNHN d’Orbigny Collection 8194, Voigt Pho-
tocard 8321).
autozooids
apertural length:
apertural width:

ca 0:21-0:25 mm
ca 0:20-0:22 mm

eleozooids (probably intramurally budded)
apertural length: ca 0-17 mm
apertural width: ca 0-13 mm

REMARKS. The description given above is based on examina-
tion of the lectotype using an optical microscope during a visit
to the MNHN in 1985. Unfortunately, the species has not
been studied using SEM and therefore detailed morphologi-
cal information is lacking (type specimens from the d’Orbigny
Collection, regrettably and much to the detriment of scientific
knowledge, are not available for loan).

Semielea dichotoma differs from S. vieilbanci in having
larger and more prominent autozooidal apertures which
occupy a high proportion of the zooidal frontal surface, and
intramurally budded eleozooids with triangular apertures.

P. D. TAYLOR

Fig. 120 Semielea vieilbanci (d’Orbigny, 1850), BMNH D4674,
Craie Marneuse, Chinon, France; gonozooid with broken frontal
wall and sediment-obscured ooeciopore, x 53.

DISTRIBUTION. Senonian (?Turonian) of the Loire Region,
France.

Genus REPTOMULTELEA @ Orbigny, 1853

TYPE SPECIES. Reptomultelea tuberosa dOrbigny, 1853: p.
655, by monotypy, subjective junior synonym of Reptelea
sarthacensis d Orbigny, 1853: p. 640.

OTHER SPECIES. Reptomultelea acclivata sp. nov., R. auris sp.
nov., R. betusora nom. nov. (for Diastopora tuberosa Reuss,
1874), R. bituberosa sp. nov., Semimultelea canui Voigt,
1924, R. convexa sp. nov., S. dixoni Lang, 1906, Meliceritites
filiozati Levinsen, 1912, R. goldfussi sp. nov., R. levinseni sp.
nov., R. matutina sp. nov., R. mitrus sp. nov., Diastopora
oceani d’Orbigny, 1850, R. parvula sp. nov., R. pegma sp.
noy., Semimultelea polytaxis Voigt, 1924, R. pseudopalpeb-
rosa sp. nov., R. reedi sp. nov., Semielea reussi Pergens,
1890, R. sarissata Gregory, 1899, R. scanica sp. nov., Clausi-
multelea tuberculata d’ Orbigny, 1853.

REVISED DIAGNOSIS. Eleid with encrusting, unilamellar or
multilamellar colony; autozooids fixed-walled; cancelli lack-
ing.

REMARKS. Reptomultelea is employed for a broad grouping
of 23 fixed-walled melicerititids with non-erect, unilamellar

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 47

1121 122

i
|

Fig. 124 Semielea dichotoma (d’Orbigny, 1853), photograph of
MNHN d’Orbigny Collection 8194, lectotype, Senonian
[?Coniacian], Tours, Indre et Loire, France; autozooids (some
with opercula) and eleozooids (?intramural), x 38.

or, more often, multilamellar colonies. The oldest known
species is R. matutina sp. nov. from the Lower Albian
Shenley Limestone of Bedfordshire, England, and the young-
est is R. scanica sp. nov. from the Lower Campanian of
Scania, Sweden. Considerably more species (12) are recorded

123

Figs 121-123 Semielea spp., photographs. 121-122, S. vieilbanci (d’Orbigny, 1850), MNHN d’Orbigny Collection 8200, Senonian,
?Villedieu, ?Vendome, syntype of Diastopora arborescens d’Orbigny, 1851; 121, profile of colony with intact base; 3-2; 122, underside of
base showing entrances to the axial lumens of four branches, Xx 4-1. 123, S. dichotoma (d’Orbigny, 1853), MNHN d’Orbigny Collection
8194, lectotype, Senonian [?Coniacian], Tours, Indre et Loire, France, x 3-7.

from the Cenomanian than from any other stage (see Fig.
127).

Table 4 is a key to species identification and should be used
in conjunction with Figures 125 and 126 which depict the
outline shapes of autozooidal and eleozooidal apertures. The
species of Reptomultelea are fairly easy to separate if the
material available is sufficient to access the character states of
the autozooids and eleozooids. Variation between the small
number of specimens available is relatively large for three of
the new species (R. convexa, R. goldfussi and R. mitrus)
recognized here. It is possible that one or more of these
species will eventually require taxonomic splitting when fur-
ther specimens become available.

Species of Reptomultelea are generally easily distinguished
from the encrusting bases of erect melicerititids because of
their large areal extent, typically multilamellar growth, regu-
larity and generally high proportion of autozooids relative to
eleozooids and kenozooids. Among species founded for
non-erect colonies of melicerititids, Reptoceritites rowei Gre-
gory, 1899 and Reptoceritites acutissima Voigt, 1963 fail to
meet these criteria and are considered to be the encrusting
bases of colonies of Meliceritites rather than species of
Reptomultelea.

A difficult problem concerns the phylogenetic relationships
between species of Reptomultelea and species of erect fixed-
walled melicerititids belonging particularly to Meliceritites
and Elea. Zooid-level characteristics, including eleozooid
morphology, can be more similar between given species of
Reptomultelea and erect species of these genera than with
other species of Reptomultelea. For example, the intramural
eleozovids of R. pseudopalpebrosa sp. nov. greatly resemble
those of Meliceritites palpebrosa Levinsen, while the pointed
eleozooids of R. tuberculata recall similar structures in M.

48 P. D. TAYLOR

OCOo 4

polytaxis parvula betusora acclivata canui matutina reedi scanica

| |

(\ ‘ 6

reussi convexa filiozati auris oceani sarthacensis pseudopalpebrosa tuberculata |

}

0.1 mm , |
levinseni pegma dixoni goldfussi bituberosa mitrus sarissata

Fig. 125 Outlines of autozooidal aperture shapes in species of Reptomultelea. Individual apertures were traced from SEM micrographs and
scaled using the mean apertural length determined for the species. Species are arranged according to apertural length.

eee |.

tuberculata parvula acclivata filiozati oceani scanica dixoni
I JI I\ )\

sarthacensis bituberosa pegma levenseni sarissata

Fig. 126 Outlines of eleozooidal aperture shapes in species of Reptomultelea. Individual apertures were traced from SEM micrographs and
scaled using the mean apertural length determined for the species. Species are arranged according to apertural length.

gothica Levinsen and several related species of Meliceritites. it demands the acquisition of erect growth several times, and
Such similarities raise the possibility that Reptomultelea is not (2) stratigraphical evidence suggests that Reptomultelea is not
monophyletic but may instead be polyphyletic or paraphyl- the most primitive melicerititid genus because the early

etic. The polyphyletic hypothesis necessitates convergent Albian appearance of Reptomultelea post-dates the late Bar-
evolution of species of Reptomultelea from various erect remian appearance of Meliceritites by a significant time
melicerititid species by parallel loss of the erect branches of period. However, a full phylogenetic analysis of the melicer-
colonies, together with enlargement of the encrusting colony ititids is needed to decide whether Reptomultelea is mono-
base, seemingly a relatively simple evolutionary option. The phyletic, polyphyletic or paraphyletic. Until this has been
paraphyletic hypothesis necessitates convergent evolution of accomplished, monophyly is assumed if only for reasons of
species of erect melicerititids from the primitive ‘genus’ nomenclatoral stability and pragmatic taxonomy.

Reptomultelea. This is a less attractive alternative because (1) Other genera established for encrusting melicerititids

‘Table 4 Key to the species of Reptomultelea. Note, because this
key places a high reliance on eleozooids, which may not be
developed in every specimen, identifications should be carefully
checked against the full descriptions. Furthermore, it is possible
that new material will reveal the presence of eleozooids in species
in which they are currently unknown.

MEN COZOOIASIPLESEME , asdqcceecce-cpooce mec asses cesbicseccaibioalseainess 2
[SIRT TES ClONSTN ee-noccbaascosedhosceece spa nnconeaccesnncacaseeanene 20
' 2. Eleozooid apertures larger than autozooids (i.e.
MES ENO ZOOS) Leer aceirarcacncae eaters acoso gaciteee eco eeae a ctestelnis ssinctes 3
Eleozooid apertures smaller than autozooids .................. 14
' 3. Autozooid apertures very wide, more than 0-20 mm .......... 4
Autozooid apertures narrower, less than 0-20 mm ............. 5
4. Autozooid apertures ogee arch-shaped, more than 0-35 mm
NOU Dee eos cere ens aiinwiten sins o' vcs sestcleine vi R. sarissata (p.94)
Autozooid apertures very slightly pointed, less than 0-30 mm
IEE. cecodacgeanee ducdgBERCeeeZeeed scpecdeuseautogeesces R. dixoni (p.69)
5. Eleozooid rostrum broad, well-rounded distally ................ 6
Eleozooid rostrum narrow, pointed or poorly-rounded
iS etl Venere ente se citae sw oid nee htsaiesiinw aatt woltinlclaps ives sesjseemelivin ss 10
6. Opercula deeply depressed distally; apertural shelf very wide;
eieozooid rostrum spatulate ...................... R. pegma (p. a)
CUR@INNIRS — ccgucodoadtadeeee noaneer cae CaoREEEE REceD ee eet ereeer Aamereeer ae
7. Zooids with a pair of prominent tubercles at either end of the
EM en Ceres aseeeroveroectrie = sala es cnasenenacett R. bituberosa (p.63)
OUINGAWISG: 2 onnns dba cactostanadcoepns HoBBbeoS Aabea see bo oe aaa scnennbAneAnE 8

8. Autozooid frontal wall short (less than 0-50 mm); aperture
occupying a high proportion of the frontal area
PRE R ee Sere eee cdots eye cts bie erciaAsiasilnsiebele ecis elo esiiei sa R. scanica (p. 2)
OUINGIRITIRE Ag aag GARR Rada be nee tan REE SAB ODEO S one COR SCO e Seco acer eE nM aneCrE
9. Eleozooid frontal wall longer than 0-85 mm
Be eee ae snccincteesstenascasdapssseuones samemsiaer R. oceani (p.80)
Eleozooid frontal wall shorter than 0-85 mm
300600¢ 80000 0G0 02 0p AGeoEe Tene eee nae ee Pep ERR ea perc ccaad R. filiozati (p.71)
10. Autozooid apertures very tall (length about 1-5 x width);
| eleozooid apertures only slightly longer than autozooid

apertures; kenozooids numerous .......... R. tuberculata (p.97)

| (COHPREIWIISS | hacgocptent AEaRE EE DerHEe SaneeEEReCEeteacceacrcercereeear eee 11
11. Eleozooid rostrum well-raised distally so that plane of aperture
is oblique to colony surface .................... R. acclivata (p.54)
(QHISORMIGO ‘cedpondbapsbodtocartese Sec cneebe secede meanen cere acemeRE Tras 12

12. Zooids very small — e.g. autozooid frontal wall width less than
0-21 mm; eleozooid frontal wall length less than 0-50 mm

Be aes ahaa ob cn caise nies Ipecdasaabansaracanccacceacoaccn Jkt JACLALHE (ote)
CO OT NGS Comer oe castes got ceo niera dss Meee je snfeloelaen Gs sei abitig ele selatelawies 13
13. Zooids medium-sized — e.g. autozooid frontal wall length less

| than 0-59 mm; eleozooid frontal wall length less than 0-90 mm

ee eae ete es cacecte cate scnahetensasssamiegs R. sarthacensis (p.49)
ASNASAT ICL SU ehs iar tacnes saesceuee ss scelenenne R. levinseni (p.74)
14. Eleozooid apertures inverted T-shaped (trifoliozooids) ..... 15
(OVINGIRIRS pebescososuedsencee baeae es anee cat oona se eeeeoeuonetee ieroncee 18

15. Autozooids commonly with small kenozooids forming ‘ears’ on
eithensideiofthelapertures --cescocceneecce- cee sesse R. auris (p. Wy

MOINS WIS CME reece cone cis eee a cuir, eo nujenouiontetcr geste sbicesdesmae’

16. Autozooid frontal wall less than 0-45 mm long .... R. ers

(p.85) _

(QUIORWSE cosacadhOedeneeteneene: Case Beerentenseeceerr ce earernmentcae 17

i. oT aperture less than 0-15 mm wide ....... R. betusora

p.59)

ca aperture more than 0-15 mm wide ...... R. convexa

p.66)

18. Eleozooid apertures D-shaped with a hood-like area
BUSI ioes foot wcien equ lenteconwaka anseeies R. pseudopalpebrosa (p. 2,
Eleozooid apertures tall and pointed ..................0.:0e0eee ee

19. Autozooid apertures less than 0:20 mm wide . R. canui (p. 5)

Autozooid apertures more than 0-20 mm wide ........ R. reussi

(p.91)

20. Autozooid apertures less than 0-15 mm wide ...... R. matutina

B77)

ROEHL WISE Soe os secese seco eee aesee ashwccecisanee nha hucussre Secweeneeene 21
21. Autozooid apertures lancet arch-shaped ...... R. mitrus (p.79)

Autozooid apertures rounded ..............0..:csecseeeeeeeeeeeeees 22

22. Autozooid apertures less than 0-21 mm long; crescentic

arrangement of opercular pseudopores .......... R. reedi (p.91)

Autozooid apertures more than 0-21 mm long; pseudopores

scattered widely over operculum ............. R. goldfussi (p.73)

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 49

include Reptelea d’Orbigny, 1853, Semimultelea d’Orbigny,
1853, Clausimultelea d’Orbigny, 1853 and Reptoceritites Gre-
gory, 1899. Lang (1906), who revised the ‘reptant eleids’,
used Reptelea for unilamellar species lacking eleozooids
(‘avicularia’), Reptoceritites for unilamellar species with eleo-
zooids, Semimultelea for bilamellar/multilamellar species
lacking eleozooids, and Reptomultelea for bilamellar/
multilamellar species with eleozooids (Clausimultelea he
regarded as belonging to the non-melicerititid family
Clausidae). The type species of Reptelea, Reptelea pulchella
d’Orbigny, 1853, is of uncertain identity but may be the base
of a colony of Elea lamellosa (d’Orbigny, 1850) (see p. 19),
that of Semimultelea, S. irregularis d’Orbigny, 1853 is also the
base of an Elea lamellosa colony (see p. 19), while that of
Reptoceritites, R. rowei Gregory, 1899, is the base of a colony
of Meliceritites dollfusi Pergens, 1890. Therefore, Reptelea,
Semimultelea and Reptoceritites have no value as genera for
the reception of non-erect melicerititid species as they all
represent basal parts of erect taxa. Lang’s (1906) scheme for
the generic division of melicerititids must be rejected. Clausi-
multelea, type species C. tuberculata d’Orbigny (see p. 99), is
merely a Reptomultelea with an above average proportion of
kenozooids.

Semielea dOrbigny, 1853, type species S. vieilbanci
d’Orbigny, 1853 (see p. 43), is retained as a genus distinct
from Reptomultelea despite the fact that cavariiform colonies
of various species of Reptomultelea (e.g. R. auris, R. convexa,
R. reussi) may closely resemble Semielea. These species of
Reptomultelea show variable orientations of zooids in the
initial layer whereas the zooids are orientated strictly parallel
to the branch axis in Semielea. Furthermore, it is probable
that cavariiform colonies of Reptomultelea grew loosely
around erect organisms like hydroids (cf. many modern
cavariiform cheilostomes). In contrast, the existence of ‘plat-
forms’ partitioning the axial tubes shows that this was not the
case in colonies of Semielea.

DISTRIBUTION. Lower Albian — Lower Campanian (Fig.
127), Europe and western Asia.
Reptomultelea sarthacensis d’Orbigny, 1853 Figs 2,
128-147

1826 Cellepora escharoides Goldfuss: 28 (partim), pl.12, figs
3b-c only.

1853 Reptelea sarthacensis d’Orbigny: 640, pl.
9-10, pl. 738, fig. 15.

1853 Reptomultelea tuberosa d’Orbigny: 655, pl. 741, figs
14-15.

1872 Diastopora oceani d’Orbigny; Reuss: 110 (partim), pl.
27, fig. 2 only.

1877 Diastopora acupunctata Novak: 99 (partim), pl. 6, fig.
1 only.

1890 Semielea sarthacensis (d’Orbigny); Pergens: 393.

1897a Semielea sarthacensis (d’Orbigny); Canu: 155, pl. 5,
fig. 10.

1897b Semielea sarthacensis (d’Orbigny); Canu: 749.

non 1899 Reptomultelea tuberosa d Orbigny; Gregory: 320,

fig. 37 [=Reptomutltelea bituberosa sp. nov., see p.
63].

1912 Meliceritites sarthacensis (d’Orbigny); Levinsen: 41,
pl. 1, figs 1-2.

21938 Reptoceritites zahdlkai Prantl: 31, pl. 2, fig. 9.

MATERIAL. Holotype: MNHN d’Orbigny Collection 6562

604, figs

(Fig. 129), Cenomanian, Le Mans, Sarthe, France; this
colony encrusts a concavity in a sponge.

Other material: MNHN d’Orbigny Colln 6589 (Fig. 128),
Cenomanian, Le Mans; presumed to be the holotype of R.
tuberosa. PSUB Goldfuss Colln 105B, Cenomanian, Essen,
Germany; presumed to be the specimen figured by Goldfuss
(1826: pl. 12, figs 3b, c) as Cellepora escharoides. SMD
un-numbered, specimen figured by Reuss (1872: pl. 27, fig. 2)
as Diastopora oceani d’Orbigny (Voigt photocard 2814),
Cenomanian, plenus Zone, Dresden-Plauen, Germany. EM
RE 551.763.31.A711, Cenomanian, Essen. VH 10432, 10472,
Lower Cenomanian, Miilheim-Broich, Westfalia, Germany.
VH 10437-8, top of Lower Cenomanian (orbignyi Zone) or
base of Middle Cenomanian (costatus Zone), Saint-Germain-
la-Campagne, Calvados, France, Breton Colln. VH 10463,
Lower Turonian, St Calais, Sarthe, France. VH 10466,
Upper Cenomanian, St Calais. VH 10510, Upper Cenoma-
nian (plenus Zone), Dresden-Plauen, Germany. BMNH
D3624 (2 specimens), D3631, Cenomanian, Essener Grtn-
sand, Essen, Westfalia, Germany. BMNH D4424 [only the
specimen on a serpulid tube], Cenomanian [plenus Zone],
Korycaner Schichten, Kamajk, Bohemia, Czechoslovakia.
BMNH D54294, Upper Cenomanian or Lower Turonian, nr
St Calais, Sarthe, France, Voigt Colln. BMNH D54304-5,
Upper Cenomanian (plenus Zone), Predboj, Bohemia,
Czechoslovakia, Voigt Colln. BMNH D58952-8, Upper Cen-
omanian (gourdoni Zone), Craie glauconieuse a Metioco-
ceras et Sciponoceras, St Calais road-cutting, Sarthe, Taylor
& Hammond Colln.

Questionably assigned: BMNH D59244-6, Chloritic Marl,
[Lower Cenomanian], St Catherine’s Point, Isle of Wight,
England.

DESCRIPTION. Colony unilamellar or multilamellar (Fig. 2),
individual layers about 0-21—0-27 mm thick and occasionally
growing free of the substratum with a transversely folded
basal lamina. Ancestrula not identified unequivocally; auto-
zooids from primary zone of astogenetic change with smaller,
more rounded apertures than those from zones of repetition.
Overgrowths (Figs 131, 141) originate by intrazooecial fission
of one or more basal zooids; pseudoancestrula an autozooid,
often surrounded by 6 daughter zooids, sometimes depressed

128

PoDITAYEOR

NVINVWONS9
NVINOYNL
NVIOVINOO
NVINVdWV9

NVINOLNVS

matutina
oceani

acclivata
bituberosa
goldfussi
mitrus

pseudopalpebrosa
parvula

pegma

reedi

sarthacensis

TEER

reussi
betusora
convexa
auris
dixoni
filiozati
levinseni
sarissata
tuberculata
canui
polytaxis
scanica

pete

“1

Fig. 127 Stratigraphical ranges of species of Reptomutltelea.

beneath general level of colony surface but occasionally
raised (possibly as a result of intramural budding); autozooids
in secondary zone of astogenetic change with smaller and
more rounded apertures than those in zone of repetition.

129

Figs 128, 129 Reptomultelea sarthacensis (d’Orbigny, 1853), photographs. 128, MNHN d’Orbigny Collection 6589, Cenomanian, Le Mans,
France; presumed to be the holotype of R. tuberosa d’Orbigny, 1853; x 3-6. 129, MNHN d’Orbigny Collection 6562, Cenomanian, Le
Mans, France; holotype of Reptelea sarthacensis d’Orbigny, 1853, showing autozooids, eleozooids and a gonozooid, x 18.

Organization fixed-walled. Zooids variably arranged, often
approximately quincuncial.

Autozooids (Fig. 130) moderately small, frontally elon-
gate, about 1-6-2 x longer than wide, often hexagonal in
outline, occasionally diamond-shaped, with a pointed distal
end; frontal wall occupying about half of the frontal surface,
‘slightly convex, with circular pseudopores; boundary wall
well-defined, salient. Aperture (Figs 132, 139) of medium
| size, on average about 1-5 x longer than wide, gothic arch-
shaped, pointed distally, attaining maximum width some-
where between the hinge line and mid-length; apertural rim
| well-developed, raised to form a tubercle-like protuberance
at the pointed distal end of the aperture; apertural shelf
narrow; hinge line with low teeth at either end of a median
bar. Operculum (Figs 133, 140) often preserved in-situ,
convex, with about 24 radially ovoidal pseudopores arranged
in a crescent close to the disto-lateral edge. Terminal dia-
phragms rarely present, located beneath level of apertural
shelf, some with scattered pseudopores and a central depres-
sion. Intramural buds not observed.

Eleozooids (Figs 142-145) abundant, scattered; moderately
large, frontal surface generally about 2-5 x longer than wide,
considerably longer and usually a little wider than the auto-
zooids; frontal wall occupying about half or less of the frontal
surface, convex with pseudopores as in the autozooids.
Aperture elongate, generally 2-3 x longer than wide, attain-
ing maximum width at or a little distally of the hinge line,
with a long, narrow rostral area sometimes a little spatulate;
rostrum generally depressed at its distal end; hinge line with a
/ wide median bar with small teeth at either end. Opercula
rarely preserved in-situ. Terminal diaphragms (Fig. 142) may
_be present. Intramurally budded eleozooids (Fig. 145) and
| 2autozooids present.
Kenozooids variable in abundance, scattered or aggre-
gated, often present immediately distal to eleozooids (Fig.
| 143) or in areas of disrupted growth (Figs 137, 146).

Gonozooids (Figs 146-147) moderately common, longitu-
dinally elongate, 1-5-2 x longer than wide, a short parallel-
: sided portion emerging from the maternal aperture and

becoming ovoidal to pear-shaped. Ooeciopore (Fig. 138)
transversely elongate, variable in width, its distal edge
indented internally by a hemiseptum. Atrial ring not
observed.

MEASUREMENTS.

autozooids

(10 zooids from BMNH D54294)

frontal length: mean = 0-48 mm; SD = 0-026 mm;
CV = 5-5; range = 0:44-0:51 mm
mean = 0-26 mm; SD = 0-019 mm;
CV = 7:2; range = 0:23-0:29 mm
mean = 0-21 mm; SD = 0-017 mm;
Q: CV = 8-0; range = 0-18-0-24 mm
mean = 0-14 mm; SD = 0-012 mm;
Q: CV = 8-6; range = 0-12-0-15 mm

(10 zooids from EM RE 551.763.31.A711)

frontal width:
apertural length:

apertural width:

- frontal length: mean = 0-51 mm; SD = 0-035 mm;
| CV = 6-9; range = 0-48-0-59 mm
_ frontal width: mean = 0-25 mm; SD = 0-018 mm;

CV = 7-1; range = 0:23-0:27 mm

mean = 0-18 mm; SD = 0-022 mm;
CV = 12-1; range = 0-15—0-23 mm
mean = 0-13 mm; SD = 0-016 mm;

apertural length:

apertural width:

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 51

CV = 12:8; range = 0-11-0-17 mm

(10 zooids from VH 10432)

frontal length: mean = 0-55 mm; SD = 0-030 mm;
CV = 5-5; range = 0:51-0:59 mm
0:27 mm; SD = 0-024 mm;
CV = 8:8; range = 0:24-0:32 mm
mean = 0-23 mm; SD = 0-017 mm;
CV = 7-3; range = 0:21-0:26 mm
mean = 0-15 mm; SD = 0-009 mm,
CV = 5-7; range = 0:14-0:17 mm

frontal width:

3
~
@
tS)
=|

apertural length:

apertural width:

eleozooids

(8 zooids from BMNH D54294)

frontal length: mean = 0-74 mm; SD = 0-054 mm;
CV = 7:3; range = 0-68-0-83 mm
mean = 0-29 mm; SD = 0-023 mm;
CV = 7-7; range = 0-26-0-:33 mm
mean = 0-49 mm; SD = 0-041 mm;
CV = 8-4; range = 0-44-0-54 mm
mean = 0:19 mm; SD = 0-013 mm;
CV = 6:5; range = 0:17-0:21 mm

(7 zooids from EM RE 551.763.31.A711)

frontal length: mean = 0-81 mm; SD = 0-029 mm;
CV = 3-6; range = 0:78-0:86 mm
mean = 0-31 mm; SD = 0-036 mm;
CV = 11-5; range = 0-27-0-38 mm
mean = 0-50 mm; SD = 0-050 mm;
CV = 10-1; range = 0-44-0-59 mm
mean = 0-17 mm; SD = 0-029 mm;
CV = 16-4; range = 0-15—0-23 mm

(8 zooids from VH 10432)

frontal length: mean = 0-70 mm; SD = 0-037 mm;
CV = 5:3; range = 0-63-0-75 mm
mean = 0-27 mm; SD = 0-017 mm;
CV = 6-6; range = 0:24-0:30 mm
mean = 0-38 mm; SD = 0-019 mm;
CV = 5-1; range = 0-36-0-41 mm
mean = 0-16 mm; SD = 0-011 mm;
CV = 7-0; range = 0-15—-0-18 mm

frontal width:
apertural length:

apertural width:

frontal width:
apertural length:

apertural width:

frontal width:
apertural length:

apertural width:

gonozooids
(1 zooid from holotype MNHN d’Orbigny Collection 6562)

frontal length: ca 1-43 mm
dilated frontal length: ca 1-30 mm
frontal width: ca 0-81 mm
ooeciopore length: ca 0-09 mm
ooeciopore width: ca 0-10 mm

(2 zooids from VH 10432)

frontal length: 1-58-1-74 mm
dilated frontal length: 1-37—1-53 mm
frontal width: 0-75—0-90 mm
ooeciopore length: 0-08 mm

ooeciopore width: 0-14-0-18 mm

REMARKS. Pergens (1890) first recognized that Reptomulte-
lea tuberosa d’Orbigny, 1853 was a junior synonym of R.
sarthacensis, the latter species having been described in the
same publication but with page priority. R. tuberosa is the
type species, by monotypy, of Reptomultelea d’Orbigny,
1853, and therefore R. sarthacensis is considered to be the
correct name for the type species of this genus. Gregory’s
(1899) R. tuberosa dOrbigny is a different species,
redescribed as R. bituberosa sp. nov. (p. 63).

52 PSD TAYEOR

134 135

Figs 130-135 Reptomultelea sarthacensis (d’Orbigny, 1853). 130-133, BMNH D54294, Upper Cenomanian or Lower Turonian, nr St Calais,
Sarthe, France; 130, autozooids and an eleozooid, x 72; 131, two depressed pseudoancestrulae surrounded by radiating autozooids, x 105;
132, autozooidal aperture, x 230; 133, autozooidal operculum, x 225. 134, VH 10463, Lower Turonian, St Calais, Sarthe, France; small
colony encrusting a shell, x 15. 135, EM RE 551.763.31.A711, Cenomanian, Essen, Germany; depressed area of colony showing
autozooids (with and without opercula) and eleozooids, x 35.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

Figs 136-141 Reptomultelea sarthacensis (d’Orbigny, 1853), VH 10432 Lower Cenomanian, Miilheim-Broich, Westfalia, Germany; 136,
autozooids (some with broken opercula) and eleozooids, x 48; 137, edge of overgrowth showing disrupted area with kenozooids and
eleozooids, X 40; 138, ooeciopore, x 137; 139, autozooidal aperture, x 200; 140, autozooidal operculum, x 250; 141, newly-developed
overgrowth, xX 21.

54 P. D. TAYLOR

145

144

143

Figs 142-145 Reptomultelea sarthacensis (d’Orbigny, 1853), variation in eleozooidal morphology. 142, VH 10432 Lower Cenomanian,
Mulheim-Broich, Westfalia, Germany; eleozooid with broad, almost parallel-sided rostrum and the remains of a terminal
diaphragm, x 100. 143, VH 10510, Upper Cenomanian (plenus Zone), Dresden-Plauen, Germany; eleozooid with narrow, slightly
spatulate rostrum resting on a distal kenozooid, x 75. 144, BMNH D54294, Upper Cenomanian or Lower Turonian, nr St Calais, Sarthe,
France; eleozooid with narrow, slightly spatulate rostrum, x 120. 145, EM RE 551.763.31.A711, Cenomanian, Essen, Germany; eleozooid

with broad, slightly spatulate rostrum and hosting an intramural eleozooid, x 92.

The type specimen of Reptoceritites zahalkai Prantl, 1938,
from the Turonian of Bohemia, was not available for study
during a visit in June 1986 to the Narodni Museum, Prague,
and Prantl’s illustration of a worn specimen is inadequate for
positive identification of the species. However, topotype
specimens (BMNH D54304-5) collected by Professor E.
Voigt are conspecific with R. sarthacensis.

One of the two specimens (SMD un-numbered; Voigt
photocard 2814) from the plenus Zone of Dresden figured by
Reuss (1872) as Diastopora oceani d’ Orbigny appears to be a
unilamellar colony of R. sarthacensis encrusting an oyster (pl.
27, fig. 2). Pergens (1890: p. 399) recognized that the D.
oceani of Reuss was not the same as d’Orbigny’s species and
proposed Semielea reussi for D. oceani sensu Reuss. S. reussi
is not placed in synonymy with R. sarthacensis because the
second of Reuss’s syntype specimens (pl. 27, fig. 3) is a
different species and is here selected as the lectotype of S.
reussi (see p. 91).

Among the material described from the Czechoslovakian
Cenomanian/Turonian as Diastopora acupunctata Novak,
1877, is at least one specimen belonging to R. sarthacensis.
This is the specimen shown in plate 6, fig. 1 of Novak (1877)
and registered in the Narodni Museum as 2437 (Voigt photo-
cards 6455 and 6458).

As here delimited, R. sarthacensis is a widely distributed
species ranging from the Lower Cenomanian to the Upper
Cenomanian or possibly Lower Turonian. Some morphomet-
ric differences exist between populations, but the unity of the
species is supported by the ubiquitous long, narrow eleozooid
rostra and gothic arch-shaped autozooid apertures with dis-

tally raised rims. Colonies from the Lower Cenomanian of
Westfalia tend to have eleozooid rostra rather broader than
those of younger populations, and an above average propor-
tion of kenozooids occurs in specimens from the Essener
Griinsand. The short but variable length of the eleozooid
rostra in material from the Lower Cenomanian of the Isle of
Wight (BMNH D59244-6) could be due to environmental
stunting or might indicate a specific distinction; therefore,
these specimens are questionably assigned to R. sarthacensis.

Reptomultelea levinseni sp. nov. (p. 74) from _ the
Coniacian/Santonian of northern France is very similar to R.
sarthacensis but has appreciably longer autozooid and eleo-
zooid frontal walls.

DISTRIBUTION. Lower Cenomanian-Upper Cenomanian or
Lower Turonian. Known from the Lower Cenomanian of
Germany, ?France and ?England; ?Middle Cenomanian of
France; Upper Cenomanian of Germany, France and
Czechoslovakia.

Reptomultelea acclivata sp. nov.

MATERIAL. Holotype: VH 10427, Lower Cenomanian,

Miuilheim-Broich, Westfalia, Germany. Paratype: VH 10502,
same horizon and locality as holotype. Other material: VH

un-numbered, several topotype colonies.

NAME. Acclivata, Latin for upward slope, with reference to
the upward-sloping eleozooid rostra.

DESCRIPTION. Colony unilamellar or multilamellar, each |

Figs 148-154 |

146

_ SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

Nn
Nn

147

Figs 146, 147 Reptomultelea sarthacensis (d’ Orbigny, 1853), gonozooids, VH 10432 Lower Cenomanian, Mulheim-Broich, Westfalia,
Germany; 146, gonozooid with broken frontal wall; note fouling tubuliporine cyclostome which has become enveloped proximally by an
overgrowth of melicerititid kenozooids, x 60; 147, two coalescent gonozooids sharing a single ooeciopore, X 70.

layer about 0-23 mm thick, often growing free of the substra-
tum and with a transversely folded basal lamina. Ancestrula

not observed. Overgrowths (Figs 148-149) presumed to origi-
‘nate by intrazooecial fission; pseudoancestrulae with
depressed apertures, small (ca 0-08—0-09 mm), more or less
equidimensional and less pointed than apertures of autozoo-
ids from zone of astogenetic repetition. Organization fixed-
walled. Zooids arranged in irregular quincunx.

Autozooids (Figs 150, 152) small, frontally elongate, about
twice as long as wide, subhexagonal with a moderately
pointed distal end; frontal wall generally occupying more
than half of frontal surface, pseudopores subcircular; bound-
ary wall poorly defined, subdued. Aperture of small size,
slightly elongate, about 1-1—1-2 x longer than wide, widest
between the hinge line and mid-length, slightly arched to
rounded distally; apertural rim well-developed distally where
it may be prolonged into a pointed projection; hinge line
bowed, detailed structure unclear; apertural shelf very nar-
row or absent. Operculum (Fig. 151) sometimes preserved
in-situ, flat centrally but with slightly convex sides, possessing
about 20 slit-like pseudopores arranged in a crescent parallel
to the disto-lateral edge; inner surface with narrow sclerites
forming a low arch across the distal edge of the operculum
where they are joined. Terminal diaphragms and intramural
buds not observed.

Eleozooids (Figs 148, 150, 152-153) numerous, scattered;

small, frontally elongate, about twice as long as wide, a little
longer and wider than the autozooids; frontal wall occupying
about half of the frontal surface, pseudopores circular and
present in about the same density as in the autozooids.
Aperture elongate, about twice as long as wide, attaining
maximum width close to the hinge line. Rostrum (Fig. 153)
long and narrow; in some eleozooids flat, but in most well
raised distally, standing above the colony surface by as much
as 0-3 mm, so that plane of aperture is inclined by up to 60° to
the colony surface, and often with a terminal hook directed
proximally. Opercula observed in situ only in overgrown
zooids; inner surface of displaced opercula seemingly with
marginal sclerites. Apparent intramurally budded eleozooids
have thickened rostra and shortened apertures.

Kenozooids occasionally present.

Gonozooids (Fig. 154) present in paratype (VH 10502),
frontally almost twice as long as wide, the bulbous distal part
being ovoidal or pear-shaped. Ooeciopore (Fig. 153) variably
transversely elongate, the ooeciostome sometimes reflexed
proximally. Atrial ring not observed.

MEASUREMENTS.

autozooids (10 zooids from holotype VH 10427)

frontal length: mean = 0-44 mm; SD = 0-038 mm;
CV = 8-7; range = 0-38-0-53 mm

frontal width: mean = 0:22 mm; SD = 0-012 mm;

56 P.D. TAYLOR

th ie Pree

Figs 148-153 Reptomultelea acclivata sp. nov. 148-151, VH 10427, holotype, Lower Cenomanian, Miilheim-Broich, Westfalia, Germany;
148, colony surface showing prominent eleozooids and overgrowths, <x 22; 149, small overgrowth, x 40; 150, autozooids and
eleozooids, < 55; 151, autozooidal operculum, x 250. 152-153, VH 10502, Lower Cenomanian, Miilheim-Broich; 152, autozooids and
eleozooids with distally-raised rostra, x 62; 153, ooeciopore and an eleozooid with long, raised rostrum, x 115.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 57

CV = 5-6; range = 0-21-0-24 mm
mean = 0-17 mm; SD = 0-011 mm;
CV = 6-7; range = 0-15-0-18 mm
mean = 0-15 mm; SD = 0-007 mm;
CV = 5-0; range = 0-14-0-15 mm

apertural length:

apertural width:

eleozooids (9 zooids from holotype VH 10427)

frontal length: mean = 0-48 mm; SD = 0-029 mm;
CV = 6-1; range = 0:44-0:53 mm
mean = 0-24 mm; SD = 0-020 mm;
CV = 8:2; range = 0-21-0-27 mm
mean = 0-28 mm; SD = 0-055 mm;
CV = 19-6; range = 0-21-0-38 mm
mean = 0-14 mm; SD = 0-016 mm;
CV = 11-5; range = 0-11-0-15 mm
[nb. apertural length measured in the plane of the aperture,
i.e. obliquely to the colony surface]

frontal width:
apertural length:

apertural width:

gonozooid (6 zooids from holotype VH 10427)

frontal length: mean = 1-50 mm; SD = 0-056 mm;
CV = 3-7; range = 1:44-1:59 mm
mean = 1-30 mm; SD = 0-044 mm;
CV = 3-4; range = 1-25-1-35 mm
mean = 0-84 mm; SD = 0-046 mm;
CV = 5-5; range = 0-80-0-90 mm
mean = 0-08 mm; SD = 0-006 mm;
CV = 7-9; range = 0-08-0-09 mm
mean = 0-14 mm; SD = 0-028 mm;
CV = 21-1; range = 0-11-0-17 mm

distal frontal wall
length
frontal width:

ooeciopore length:

ooeciopore width:

REMARKS. This new resembles Reptomultelea

species

Fig. 154 Reptomutltelea acclivata sp. nov., VH 10502, Lower
Cenomanian, Miilheim-Broich, Westfalia, Germany,
gonozooid, x 70.

sarthacensis (d’Orbigny) and R. parvula sp. nov., with which
it co-occurs at Mtilheim. However, it differs in having abun-
dant eleozooids with distally raised rostra, causing the plane
of the aperture to be at a considerable angle to the colony
surface and the colony to have a spiny appearance. In
well-preserved specimens the distal ends of the rostra are
slightly hooked, a feature unique to R. acclivata among
melicerititids. Autozooid apertures are less elongate than in
R. sarthacensis and larger than in R. parvula.

The holotype colony evidently encrusted a fragment of a
dendroid cyclostome, grew free beyond its initial substra-
tum (as with many other colonies of Reptomultelea spp.
from Miilheim), and incorporated secondary substrates,
including an onychocellid cheilostome bryozoan, into its
base.

DISTRIBUTION. Lower Cenomanian of Mtilheim, Westfalia,
Germany.

Reptomuitelea auris sp. nov. Figs 155-162
MATERIAL. Holotype: BMNH D46049, Turonian, lata Zone,
Ballard Point, Dorset, England, A.W. Rowe Collection.

Paratypes: BMNH D46048, D46052, same details as holo-
type. D43694, [Coniacian], M. cortestudinarium Zone,
Seaford Head, Sussex, England, A.W. Rowe Colln. D46027,
Turonian, planus Zone, White Nothe, Dorset, England,
A.W. Rowe Colln. BZ 1005-6, Turonian, planus Zone
Chalk, Tilleul Beach, Seine Maritime, France, Taylor and
Hammond Colln, 1985. BZ 1007, Coniacian, Craie 4 M.
normanniae, above Tilleul No. 3 Hardground (see Kennedy
and Juignet, 1974), Etretat, Seine Maritime.

NAME. Auris, Latin for ear, with reference to the paired
kenozooids which resemble ears on either side of the auto-
zooidal aperture.

DESCRIPTION. Colony unilamellar or multilamellar, each
layer about 0-3 mm thick, generally (?always) cavariiform
with tubular branches 5—10 mm in diameter, up to 40 mm in
maximum observed length, flexuous and occasionally bifur-
cating; basal lamina on inside of tubes with transverse undu-
lations, apparently growing freely of a substratum.
Overgrowths (Fig. 158) originate by intrazooecial fission;
pseudoancestrula an autozooid, aperture depressed, small,
about 0-11-0-12 mm long by 0-08 mm wide, surrounded by
approximately 6 periancestrular buds initiating a secondary
zone of astogenetic change of increasing zooid size. Organiza-
tion fixed-walled. Zooidal apertures arranged roughly in
quincunx away from overgrowth origins and anastomoses.
Autozooids (Figs 155-156, 159, 161) of large size, frontally
elongate, usually a little over twice as long as wide, subhex-
agonal or subrhomboidal in outline, subacuminate distally;
frontal wall convex, pseudopores circular; boundary wall
salient. Aperture (Fig. 160) of moderate size, longitudinally
elongate, on average 1-1-1-2 x longer than wide, attaining
maximum width just distal to the hinge line, arched distally;
apertural rim narrow; apertural shelf moderately wide dis-
tally, tapering proximally; hinge line bowed, raised slightly so
that plane of aperture faces distally. Operculum (Figs 157,
162) often preserved in-situ, convex; pseudopores slightly
radially elongate, numbering about 20, arranged close to and
parallel with the distal edge of the operculum. Terminal

a
157

P. D. TAYLOR

158

Figs 155-158 Reptomultelea auris sp. nov., BMNH D46049, holotype, Turonian, /ata Zone, Ballard Point, Dorset, England; 155,
autozooids, eleozooids and kenozooids, x 35; 156, operculate autozooids, kenozooids, intramural eleozooid (left), and evidence of
regeneration after damage in upper left autozooid, x 72; 157, autozooidal operculum, Xx 240; 158, overgrowth origin, x 56.

diaphragms not observed. Intramurally budded eleozooids
(Fig. 156) common, apertures trifoliate like those of primary
eleozooids, elevated distally so that plane of aperture faces
proximally.

Kenozooids (Figs 155-156) abundant, small, commonly
paired on either side of the autozooidal apertures and semi-
circular or crescent-shaped in outline.

Eleozooids (Fig. 161) infrequent, scattered, frontally elon-
gate, typically slightly longer but narrower than the autozoo-
ids, narrow and pointed distally. Aperture small,
longitudinally elongate, trifoliate in outline, about twice as
long as wide, considerably shorter and narrower than auto-
zooidal apertures; apertural rim moderately raised. Opercula
not observed. Intramural buds unknown.

Gonozooids known from a single broken example which is

missing the proximal part; frontal shape apparently subcircu-
lar. Ooeciopore transversely elliptical, slightly more than
2 X wider than long. Atrial ring present.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from holotype
BMNH D46049)
frontal length: mean = 0-66 mm; SD = 0-069 mm;
CV = 10-4; range = 0-54-0-78 mm
mean = 0:28 mm; SD = 0-025 mm;
CV = 8-7; range = 0-26-0-33 mm
mean = 0-20 mm; SD = 0-008 mm;
CV = 3-8; range = 0-20-0-21 mm
mean = 0-17 mm; SD = 0-011 mm;
CV = 6-1; range = 0-15-0-18 mm

frontal width:
apertural length:

apertural width:

|
|

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 59

eleozooids (3 zooids from holotype BMNH D46049)
| frontal length: range = 0-65-0-87 mm
frontal width: range = 0-24-0-29 mm
apertural length: range = 0-12-0-14 mm

apertural width: range = 0-06—0-08 mm

gonozooid (1 incomplete zooid from BMNH BZ1006)

frontal width: ca 1-13 mm
| ooeciopore length: 0-09 mm
| ooeciopore width: 0-20 mm

| REMARKS. Zooidal dimensions in this new species are similar
to Reptomultelea canui (Voigt), but R. auris differs in having
autozooidal apertures more pointed distally and inclined
‘slightly in a distal direction, and also in the presence of
common small kenozooids on either side of the autozooidal
apertures. These ‘ear-like’ paired kenozooids are not ubiqui-
‘tous but are associated with a significant proportion of

autozooids in all colonies of R. auris. They provide the most

Figs 159-162 Reptomultelea auris sp. nov., 159, BZ 1005, Turonian, planus Zone Chalk, Tilleul Beach, Seine Maritime, France, operculate
autozooids with small kenozooids lateral to the opercula, x 82. 160-162, BMNH D46049, holotype, Turonian, /ata Zone, Ballard Point,
Dorset, England; 160, autozooidal aperture, x 182; 161, autozooids and primary eleozooid, x 65; 162, autozooidal operculum bordered
distally and laterally by a kenozooid, x 125.

useful means of recognizing the species, being known only
from this species among melicerititids.

The cavariiform morphology (sensu Brood, 1972) of R.
auris colonies with free-growing basal laminae suggests loose
growth of colonies around, but not directly encrusting,
arborescent organisms. Modern colonies of Schizoporella and
other cheilostomes show a similar morphology when growing
loosely around clusters of hydroid stems or other erect
substrates.

DISTRIBUTION. Upper Turonian-Coniacian of southern

England and northern France.

Figs 163-173

21846 Diastopora confluens Reuss: 65, pl. 15, figs 41, 42.
1874 Diastopora tuberosa Reuss: 132, pl. II. 25, figs 2 and
3h:
1877 Diastopora acupunctata Novak: 99 (partim), pl. 6,

Reptomultelea betusora nom. nov.

167 | callie 7 AR

Figs 163-168 Reptomultelea betusora nom. nov. 163-164, SMD un-numbered, lectotype, Turonian, Strehlen, Dresden, Germany; 163,
pseudoancestrula surrounded by radiating autozooids, x 80; 164, oblique view showing four eleozooids, x 100. 165-167, VH 10436, Upper

Cenomanian, plenus Zone, Kank, Kutna Hora, Czechoslovakia; 165, autozooids and intramural eleozooids, x 100; 166,
ooeciopore, X 180; 167, general view of colony with two broken gonozooids, x 21. 168, BMNH BZ 1000 Coniacian, Craie 4 Micraster

normanniae, above Tilleul No. 3 Hardground, Etretat, Seine-Maritime, France, part of large colony, X 22.

figs 2-5, ?figs 6-14 [non fig. 1 = R. sarthacensis].

21877 Diastopora acupunctata Novak; Fric: 93, 146, fig.
149.

21883 Diastopora acupunctata Novak; Fric: 81, 124, 125,
fig. 104.

21892 Semielea acupunctata (Novak); Pocta: 29, pl. 2, fig.
16.

Semimultelea acupunctata (Novak); Gregory: 297.

MATERIAL. Lectotype: SMD un-numbered, the specimen
figured here as Figs 163, 164, Turonian, Strehlen, Dresden,
Germany. Paralectotypes: SMD un-numbered, 2 specimens
in same sample as lectotype.

| Other material: VH 10436, Upper Cenomanian, plenus
Zone, Kank, Kutna Hora, Czechoslovakia. BMNH BZ 999,
Cenomanian [plenus Zone], Korycaner Schichten, Kamajk,
Bohemia, Czechoslovakia, Fric Colln. D58949-50, Turonian,
nodosoides Zone, Craie a Inoceramus labiatus, St Calais
road-cutting, Sarthe, France, Taylor & Hammond Colln
1985. BZ 1000-4, Coniacian, Craie 4 Micraster normanniae,
above Tilleul No. 3 Hardground (see Kennedy & Juignet,
1974), Etretat, Seine-Maritime, France, Taylor & Hammond
Colin 1985.

-
NAME. An anagram of tuberosa.

DEscRIPTION . Colony unilamellar or multilamellar, each
jayer about 0-35—-0-40 mm thick, sometimes growing freely
and with a transversely wrinkled basal lamina. Ancestrula not
bserved. Overgrowths originate by intrazooecial fission;
seudoancestrula (Fig. 163) usually an autozooid (aperture ca
‘10 X 0-08 mm diameter), sometimes an eleozooid, sur-
‘ounded by about 6 daughter buds which initiate a secondary
zone of astogenetic change of increasing zooid size. Organiza-
sion fixed-walled. Zooidal apertures variably arranged, often
-n irregular quincunx.

i Autozooids (Figs 165, 169) medium-sized, frontally elon-
J

ate, On average about twice as long as wide, often
“jexagonal or rhomboidal in outline, rounded distally.
Frontal wall slightly convex, occupying about two-thirds of
the frontal surface, slightly convex; boundary wall salient.
Aperture (Fig. 172) small, elongate, about 1:2-1-:5 x
jonger than wide, attaining maximum width about mid-
jength, moderately rounded distally; apertural rim raised
‘plightly, especially distally; apertural shelf narrow, taper-
‘ng proximally. Operculum (Fig. 170) seldom preserved
‘jn-situ, convex with a crescent of elongate pseudopores.
Terminal diaphragms not observed. Intramurally budded
-2leozooids (Figs 165, 171) frequent, intramural autozooids
1ot seen.

Kenozooids present in varying numbers, sometimes abun-
ant and clustered.

Eleozooids (Figs 164, 169, 171) common; frontal surface
enerally a little smaller than autozooids but similar in
verall proportions, although much narrower distally in the
pertural region. Aperture elongate, on average about
twice as long as wide, considerably smaller than an auto-
zooidal aperture, inverted T-shaped in outline because of
indentation by rostral shelf, often set in a plane oblique to
the colony surface and Uirected proximally. Operculum not
observed in-situ.

_ Gonozooids (Figs 167, 173) known only from specimen VH

0436, which has two, both budded close to the origin of an

vergrowth. Frontally relatively small, elongate, about twice

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 61

as long as wide, the distal inflated frontal wall emerging from
the maternal aperture, initially parallel-sided before dilating
and becoming ovoidal. Ooeciopore (Fig. 166) transversely
elongate, indented by a proximal hemiseptum. Atrial ring
present. Floor of gonozooid has salient outlines of underlying
zooids beneath proximal part of dilated frontal wall,
smoother distally.

MEASUREMENTS.

autozooids

(10 zooids from VH 10436)

frontal length: mean = 0-45 mm; SD = 0-042 mm;
CV = 9-5; range = 0:39-0:53 mm
mean = 0-21 mm; SD = 0-019 mm;
CV = 8-8; range = 0-20-0-24 mm
mean = 0:14 mm; SD = 0-009 mm;
CV = 6-9; range = 0-12-0-15 mm
mean = 0-11 mm; SD = 0-008 mm;
CV = 7-0; range = 0-11-0-12 mm

(10 zooids from BMNH BZ 1000)

frontal length: mean = 0-54 mm; SD = 0-040 mm;
CV = 7-4; range = 0:50-0:63 mm
mean = 0-28 mm; SD = 0-020 mm;
CV = 7-0; range = 0-26-0-32 mm
mean = 0-16 mm; SD = 0-014 mm;
CV = 9-1; range = 0:14-0:18 mm
mean = 0-12 mm; SD = 0-009 mm;
CV = 7-0; range = 0:11-0:14 mm

frontal width:
apertural length:

apertural width:

frontal width:
apertural length:
apertural width:

eleozooids

(VH 10436)
frontal length:
frontal width:
apertural length:
apertural width:

ca 0:35-0:38 mm
ca 0:18-0:20 mm
ca 0:06-0:08 mm
ca 0:05 mm

(5 zooids from BMNH D59329)

frontal length: mean = 0-50 mm; SD = 0-048 mm;
CV = 9-6; range = 0-42-0-54 mm
mean = 0-26 mm; SD = 0-017 mm;
CV = 6-6; range = 0:24-0:29 mm
mean = 0-09 mm; SD = 0-024 mm;
CV = 26-4; range = 0-06-0-12 mm
mean = 0-04 mm; SD = 0-008 mm;
CV = 21-1; range = 0-03—0-05 mm
gonozooids (2 zooids from VH 10436)

frontal width:
apertural length:

apertural width:

frontal length: 1-20-1-29 mm

distal frontal wall 0-99-1-11 mm
length:

frontal width: 0-53—0-68 mm

ooeciopore length: 0-06 mm

ooeciopore width: 0-09-0-11 mm

REMARKS. Reassignment of the Reuss (1874) species tube-
rosa from Diastopora to Reptomultelea makes it a secondary
junior homonym of Reptomultelea tuberosa d’Orbigny, 1853.
Although the latter species, which is the type species of
Reptomultelea, is a subjective junior synomym of R.
sarthacensis d’Orbigny, 1853 (see p. 51), a replacement name
is nevertheless required for the Reuss species (P.K. Tubbs,
ICZN, pers comm. September 1990). This not only avoids
potential problems should the synonymy between R. tuberosa
and R. sarthacensis be rejected sometime in the future, but
also prevents possible confusion between the species of Reuss
and the name of the valid type species of Reptomutltelea.

P. D: TAYLOR

Figs 169-172 Reptomultelea betusora nom. noy., BMNH BZ 1000 Coniacian, Craie 4 Micraster normanniae, above Tilleul No. 3 Hardground,
Etretat, Seine-Maritime, France; 169, autozooids and eleozooids, x 46; 170, autozooidal operculum, x 270; 171, apertures of primary (top
left) and intramural (lower centre) eleozooids, x 160; 172, autozooidal aperture, x 225.

Therefore, betusora is here proposed as a nom. nov. for the
tuberosa of Reuss (1874).

The types of Diastopora acupunctata Novak, 1877 were not
available for study during a visit to the Narodni Museum,
Prague during June 1986- However, judging from Novak’s
figures and Voigt photocards of this material, some of
Novak’s specimens belong to Reptomultelea betusora, at least
one to Reptomultelea sarthacensis d’Orbigny, and several
others cannot be identified with any reasonable confidence.

Diastopora confluens Reuss, 1846 (non Roemer) is placed
tentatively in the synonymy of R. betusora. According to
Prof. E. Voigt (pers comm., February 1987), the type speci-
men was destroyed in 1956, but a similar specimen exists in
the collections of the Naturhistorisches Museum, Vienna
(Voigt photocard 8650).

Reptomultelea betusora is characterized by the presence of
small eleozooids of the trifoliozooid type. In this respect it

resembles R. tuberculata (d’Orbigny, 1853) but the latter }

species has more elongate autozooidal and eleozooidal aper-
tures.

Specimens of R. betusora from the Turonian of Sarthe and
from the Coniacian of Etretat have rather larger zooids than
Upper Cenomanian material; however, the difference is not
considered sufficient to warrant species separation. Colonies
are often nodular in form, like those of R. /evinseni sp. nov.
(see p. 77) and R. sarissata Gregory (p. 94) which also occur |
in bryozoan-rich chalks of the Seine Maritime.

DISTRIBUTION. Upper Cenomanian (plenus Zone) to Conia-
cian, Czechoslovakia, Germany and France.

ea ay. a ee eee gee eee cee

ig. 173 Reptomultelea betusora nom. nov., VH 10436, Upper
Cenomanian, plenus Zone, Kank, Kutna Hora, Czechoslovakia,
gonozooid with broken frontal wall, x 75.

Figs 174-180

1899 Reptomultelea tuberosa d’Orbigny; Gregory: 320, fig.
ile

eptelea bituberosa sp. nov.

ATERIAL. Holotype: BMNH 36746, ‘Albian, Upper
Greensand’, Ventnor, Isle of Wight, England, Norman Col-
ection. This specimen comprises one large piece (70 x 45
mm) in matrix, a coated stub with several small fragments,
and 7 fragments mounted in two cavity slides. Although the
orizon is given as Albian, the close similarity of the matrix to
that of a better localized paratype specimen (D58206) from
ithe Isle of Wight suggests that the specimen almost certainly
‘comes from the Lower Cenomanian (carcitanense Zone)
‘Glauconitic Marl (formerly known as the Chloritic Marl).

Paratypes: BMNH D44610, ‘Upper Greensand’ [? Glauco-
‘nitic Marl], Niton, Isle of Wight, Ford Colln. BMNH D58206
(2 fragments), Lower Cenomanian, carcitanense Zone, basal
}conglomerate of the Glauconitic Marl, Rocken End, Isle of
Wight, A. Gale Colln. BMNH D59207, ‘Chloritic Marl’, St

(Catherine's Point, Isle of Wight.

‘NAME. With reference to the two prominent tubercles at
jeither end of the hinge line.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 63

DESCRIPTION. Colony encrusting, multilamellar, characteris-
tically large with up to at least 35 layers, each layer about 0-25
mm in thickness. Overgrowths (Fig. 179) develop by eruptive
budding onto the colony surface, often from several closely-
spaced pseudoancestrulae; pseudoancestrula aperture about
0-14 mm long by 0-11 mm wide; peri-pseudoancestrular
zooids usually autozooids, occasionally eleozooids. Ances-
trula not observed. Organization fixed-walled.

Autozooids (Fig. 174) of very large size, frontally elon-
gate, 2-3 x longer than wide, often hexagonal in outline
shape, rather elongate and pointed distally; frontal wall
occupying about half of the frontal surface, flat or slightly
convex with circular pseudopores; zooecial boundary wall
moderately raised. Aperture (Fig. 176) large, very longitu-
dinally elongate, about twice as long as wide, attaining
maximum width approximately mid-length, moderately
rounded distally; apertural rim well-developed, proximally
continuous with prominent tubercles paired at either end of
the hinge line; apertural shelf broad distally, tapering
proximally towards the hinge line; hinge line raised so that
plane of aperture slopes downwards in a distal direction,
with a short median bar running between two hinge teeth.
Operculum (Fig. 175) often preserved in-situ, convex, with
a crescent of slit-shaped pseudopores; sclerite impressions
visible on sediment/cement infilling zooids. Terminal dia-
phragms (Fig. 178) sometimes present, flat, located just
beneath level of apertural rim. Intramurally budded auto-
zooids (Fig. 177) observed but uncommon; in-situ opercula
may occur.

Eleozooids (Figs 179, 180) abundant, scattered; frontal
surface about 2-3 x longer than wide, significantly longer
and a little wider than an autozooid; frontal wall occupying
less than half of the frontal surface, with circular pseudopores
present in a similar density to an autozooid. Aperture elon-
gate, over twice as long as wide, widest at the hinge line,
initially tapering rapidly and then becoming parallel-sided or
slightly spatulate, well-rounded distally; rostrum forming an
extensive platform extending beyond distal end of opercu-
lum; apertural rim less prominent than that of an autozooid,
tubercles absent; hinge line with a wide median bar. Opercula
(Fig. 180) often preserved in-situ, surface convex with a
crescent of slit-shaped pseudopores; distal end of closed
operculum depressed well beneath level of apertural rim;
sclerite impressions visible on sediment/cement infilling zoo-
ids. Intramurally budded eleozooids and autozooids
observed.

Kenozooids (Fig. 174) developed in moderate numbers,
scattered among the autozooids.

Gonozooids present only in the holotype which has two
examples, both occurring in areas of overgrowth anastomosis
and distorted. Frontal wall about twice as long as wide, with a
parallel-sided portion emerging from the maternal aperture,
pear-shaped distally. Ooeciopore poorly-preserved in one
gonozooid, absent in the second.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from holotype
BMNH 36746)
frontal length: mean = (0.78 mm; SD = 0-069 mm;
CV = 8-9; range = 0-65-0-89 mm
mean = 0-32 mm; SD = 0-038 mm;
CV = 12:1; range = 0-26-0-38 mm
mean = 0-31 mm; SD = 0-016 mm;
CV = 5-1; range = 0-29-0-35 mm

frontal width:

apertural length:

178

Figs 174-179 Reptomultelea bituberosa sp. nov., BMNH 36746, holotype, ‘Albian, Upper Greensand’ [probably Lower Cenomanian
Glauconitic Marl], Ventnor, Isle of Wight, England; 174, autozooids and kenozooids, x 35; 175, autozooidal operculum, x 150; 1
autozooidal aperture with sediment mould of inner surface of operculum, x 150; 177, operculum of intramural autozooid, x 150; 17

autozooidal aperture closed by pseudoporous terminal diaphragm, x 150; 179, overgrowth origin including peri-pseudoancestrular
eleozooid, x 50.

mean = 0-16 mm; SD = 0-010 mm;
CV = 6-7; range = 0-15-0-18 mm

eleozooids (8 zooids from holotype BMNH 36746)
frontal length: mean = 0:92 mm; SD = 0-065 mm;
| CV = 7-0; range = 0-83-1-00 mm
frontal width: mean = 0-36 mm; SD = 0-060 mm;
CV = 16-7; range = 0:30-0:50 mm
mean = 0:54 mm; SD = 0-044 mm;
CV = 8-1; range = 0-48-0-60 mm
pectnral width: mean = 0-24 mm; SD = 0-029 mm;
CV = 12-0; range = 0-21-0-29 mm

gonozooid (one zooid from holotype BMNH 36746)

| apertural width:

| apertural length:

frontal length: 2-61 mm

distal frontal wall 2:34 mm
length:

frontal wall width: 1-28 mm

REMARKS. Gregory (1899) incorrectly attributed the holo-
type specimen of this new species to Reptomultelea tuberosa
d’Orbigny, 1853, a subjective junior synonym of Reptomulte-
lea sarthacensis (d’Orbigny, 1851) which is distinguished

Fig. 180 Reptomultelea bituberosa sp. nov., BMNH 36746, holotype,
‘Albian, Upper Greensand’ [probably Lower Cenomanian
Glauconitic Marl], Ventnor, Isle of Wight, England; secondary
electron image of coated specimen showing eleozooid with in-situ
operculum, x 130.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 65

by the more acute eleozooids (see p. 51). Furthermore, R.
bituberosa differs from R. sarthacensis and other species of
Reptomultelea in having prominent tubercles at either end of
the hinge line in autozooids, and autozooid apertures which
are extremely elongate, about twice as long as wide.

DISTRIBUTION. Lower Cenomanian, Glauconitic Marl (carci-
tanense Zone) of the Isle of Wight.

Reptomultelea canui (Voigt, 1924)
185-190

1924 Semimultelea canui Voigt: 167, pl. 4, figs 12-13.

Figs 181-183,

MATERIAL. Type: Although the type material of this species
was destroyed during World War 2, the unequivocal identity
of the species means that it is not necessary to select a
neotype.

Other material: BMNH D39514, D39518 (2 pieces), Santo-
nian, Gr. Bilten, Peine, Westfalia, Germany. VH 10305,
Lower Santonian, Vallstedt, Westfalia, Germany.

DESCRIPTION. Colony multilamellar, each layer about 0-5
mm thick, the basal surface of the colony sometimes growing
free from the original substrate of encrustation and incorpo-
rating secondary substrates (Figs 181-183). Overgrowth ori-
gins depressed; pseudoancestrula an autozooid, an eleozooid
or a kenozooid, surrounded by 6~7 radiating buds (presum-
ably produced by intrazooecial fission) initiating a secondary
zone of astogenetic change of increasing zooid size. Organiza-
tion fixed-walled. Zooidal apertures arranged in quincunx
except where disrupted near overgrowth origins and anasto-
moses.

Autozooids (Figs 185, 188) of medium size, frontally
elongate, almost twice as long as wide, rhomboidal in outline,
well-rounded distally; frontal wall gently convex,
pseudopores subcircular; boundary wall salient. Aperture
(Fig. 187) small, usually longitudinally elongate, up to 1-2 x
longer than wide, attaining maximum width about mid-
length, well-rounded distally; apertural rim raised, continu-
ous with zooidal boundary wall, thickened and prominent at
proximo-lateral corners of apertures; apertural shelf narrow;
hinge line bowed. Operculum (Fig. 186) often preserved
in-situ (especially in older zooids exposed by exfoliating an
overlying layer of zooids), convex; pseudopores radially
elongate, arranged in an irregular crescent and numbering
about 18-20- Terminal diaphragms (Fig. 187) often present at
a level just proximal to the apertural shelf, pseudopores not
evident (?absent) in most, abundant in some. Intramurally
budded eleozooids common, apertures (Fig. 190) of very
similar size and shape to primary eleozooids; intramurally
budded autozooids not observed.

Kenozooids infrequent.

Eleozooids (Figs 185, 188) moderately abundant, frontally
elongate, about the same length and width as the autozooids
but more pointed distally. Aperture (Fig. 189) elongate,
about twice as long as wide, narrow arch-shaped, attaining
maximum width about mid-length; apertural rim prominent
distally. Opercula not observed in-situ. Intramural buds not
seen.

Gonozooids unknown.

MEASUREMENTS.

autozooids (10 zooids from VH 10305)

frontal length: mean = 0-58 mm; SD = 0-054 mm;
CV = 9-4; range = 0-51-0-71 mm

P. D. TAYLOR

181

183

182

fel |
* *

*
Treete

tie

o>
a)

184

Figs 181-184 Reptomultelea spp., photographs of colonies. 181-183, R. canui (Voigt, 1924), VH 10305, Lower Santonian, Vallstedt,
Westfalia, Germany, < 2-9; 181, upper side of the colony showing chimney-like growth around an unpreserved object; 182, profile; 183,
underside showing original substrate (centre), two secondary substrates (lower right), and free colony growth. 184, R. goldfussi sp. nov.,
EM RE 551.763.31.A745/1, holotype, Cenomanian, Essen, Westfalia, Germany, worn upper surface of colony, X 3-3.

mean = 0:30 mm; SD = 0-023 mm;
CV = 7-8; range = 0:27-0:32 mm
mean = 0-17 mm; SD = 0-010 mm;
CV = 6-0; range = 0-15-0-18 mm
mean = ():15 mm; SD = 0-007 mm;
CV = 4-7; range = 0-15-0-17 mm

eleozooids (5 zooids from VH 10305)

frontal length: range = 0-56-0-60 mm
frontal width: range = 0-30-0-33 mm
apertural length: range = 0-12-0-15 mm
apertural width: range = 0-06—0-08 mm

frontal width:
apertural length:

apertural width:

REMARKS. Reptomultelea canui co-occurs with the com-
moner R. polytaxis (Voigt, 1924) in the Santonian of Westfa-
lia and can be distinguished from the latter by its larger

autozooids and less numerous kenozooids. The combination
of well-rounded autozooidal apertures and narrow arch-
shaped eleozooidal apertures can be used to recognize R.
canui among other species of Reptomutltelea.

DISTRIBUTION. Santonian of Westfalia, Germany.

Reptomultelea convexa sp. nov. Figs 191-196

MATERIAL. Holotype: VH 10467, Turonian (?), Chenu,
between Le Mans and Tours, Sarthe, France, collected by G.
Breton.

Paratypes: VH 10443, 10444, 10550 (8 fragments), Turo-
nian, Bois de Gareau, near Ecommoi, Sarthe, France.

TEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

Figs 185-190 Reptomultelea canui (Voigt, 1924), VH 10305, Lower Santonian, Vallstedt, Westfalia, Germany; 185, autozooids and
eleozooids, x 40; 186, autozooidal operculum, x 300; 187, autozooidal aperture closed by terminal diaphragm, x 250; 188, autozooids and
eleozooids, X 85; 189, primary eleozooid aperture, X 250; 190, intramural eleozooid aperture, x 250.

Figs 191-196 Reptomultelea convexa sp. nov. 191-193, VH 10467, holotype, Turonian (?), Chenu, between Le Mans and Tours, Sarthe,
France; 191, autozooids and eleozooids, x 42; 192, autozooidal operculum, x 200; 193, autozooidal aperture with broken terminal
diaphragm, x 200. 194-196, Turonian, Bois de Gareau, near Ecommoi, Sarthe, France. 194, VH 10444, partly-formed, broken gonozooid
(upper left) and overgrowth (upper right), x 33. 195-196, VH 10443; 195, intramural eleozooids and autozooids, X 95; 196, aperture of
intramural eleozooid, Xx 200.

NAME. With reference to the marked convexity of the distal
part of the frontal wall.

DESCRIPTION. Colony unilamellar or multilamellar, each
layer between 0-23 and 0-30 mm thick, often cavariiform and
generally with a free-growing, undulose basal lamina. Over-
‘growths (Fig. 194) originate through intrazooecial fission.
Organization fixed-walled. Apertures arranged roughly in
-quincunx.

Autozooids (Figs 191, 195) medium-sized, frontally elon-
igate, less than twice as long as wide, usually subhexagonal or
subrhomboidal in outline, subrounded distally; frontal wall
iconvex, especially close to the hinge line; boundary wall
‘salient. Aperture (Fig. 193) of moderate size, occupying
about a third of the frontal surface, longitudinally elongate,
sabout 1-2-1-3 x longer than wide, arched distally, attaining
‘maximum width about mid-length (or a little proximally of
‘mid-length); apertural rim slightly raised; hinge line poorly
preserved in all available specimens; apertural shelf conspicu-
‘ous, wide to moderately wide, tapering proximally. Opercu-
lum (Fig. 192) convex; pseudopores not visible in poorly
‘preserved specimens. Intramurally budded eleozooids com-
‘mon; aperture often cowl-like, directed proximally (Fig. 190).

Kenozooids infrequent.

Eleozooids (Figs 185, 188) common, often clustered in
groups of 2 or 3, frontally elongate, less than twice as long as
wide and smaller than the autozooids, pointed distally. Aper-
jture (Fig. 189) of small size, trifoliate, about twice as long as
wide. Opercula not observed. Intramurally budded eleozoo-
lids apparently present, distinguished from ‘non-regenerated’
‘eleozooids by proximal inclination of apertural plane.
Gonozooids known from one complete and one broken
example (Fig. 194). Frontally of small size, elongate, over
twice as long as wide, a parallel-sided tube of variable length
emerging from the maternal aperture and dilating into a
subcircular distal portion. Ooeciopore transversely elongate,

|twice as wide as long. Atrial ring not observed.

MEASUREMENTS.

autozooids (10 zooids from holotype VH 10467)

frontal length: mean = 0:55 mm; SD = 0-053 mm;
CV = 9-6; range = 0-48-0-62 mm
mean = 0-33 mm; SD = 0-017 mm;
CV = 5-0; range = 0:30-0:35 mm
mean = 0-20 mm; SD = 0-009 mm;
CV = 4:3; range = 0-18-0-21 mm
mean = 0-16 mm; SD = 0-007 mm;
CV = 4-5; range = 0:15-0:17 mm

{

apertural length:

apertural width:

mean = 0:49 mm; SD = 0-052 mm;
CV = 10-6; range = 0-44-0-57 mm
mean = 0-28 mm; SD = 0-035 mm;
CV = 12-4; range = 0:21-0:32 mm
mean = 0-12 mm; SD = 0-019 mm;
CV = 16-6; range = 0-09-0-14 mm
mean = 0:06 mm; SD = 0-005 mm;
CV = 8-6; range = 0-06-0-08 mm

gonozooids (1 zooid from VH 10550)

frontal length 1-62 mm

distal frontal wall 1-26 mm
length:

frontal width: 0-71 mm
oeciopore length: 0-08 mm

ooeciopore width: 0-15 mm

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 69

REMARKS. This species is represented only by fragmentary
specimens which are either cavariiform or free-growing
lamellar. The high convexity of the distal frontal wall close to
the aperture is particularly characteristic. Eleozooids have
trifoliate apertures like those of Reptomultelea betusora nom.
nov., R. polytaxis (Voigt) and R. auris sp. noy., but R.
convexa has much larger autozooids than R. polytaxis and is
distinguished by its convex frontal walls from all three
species.

The intramurally budded eleozooids in the paratypes (Figs
194-196) are more strongly cowled, autozooidal apertures
larger and their shelves broader than in the holotype (Figs
191-193). These differences are not considered to warrant
species level separation, however, until sufficient material is
available to assess species variability.

DISTRIBUTION. Turonian of Sarthe, France.

Reptomultelea dixoni (Lang, 1906) Figs 197-201

1899 Reptomultisparsa rowei Gregory: 121 (partim) [non pl.
7, fig. 1].
1906 Semimultelea dixoni Lang: 64, figs 4 and 12.

MATERIAL. Holotype: BMNH D7845, [Coniacian], cortestu-
dinarium Zone, Pit No. 32 of Dibley (1900), opposite the
Rose and Crown Inn, Kenley, S. of Croydon, Surrey,
England, Withers and Chatwin Collection.

Other material: BMNH D3031 (2 fragments), Upper
Chalk, Offham Pit, Lewes, Sussex, Capron Colln (originally
described as Reptomultisparsa rowei Gregory, 1899). BMNH
D31844—50 (fragments of one colony), [Turonian], Dunton
Green, Sevenoaks, Kent, England, Dibley Colln. BMNH
D43660, [Coniacian], cortestudinarium Zone, Seaford Head,
Sussex, England, Rowe Colln. BMNH D46061, D46066,
Turonian, /ata Zone, White Cliff to the Hooken, Devon,
Rowe Colln. BMNH D57505-9, Coniacian, Craie 4 M.
normanniae, above Tilleul No. 3 Hardground (see Kennedy
& Juignet, 1974), Etretat, Seine Maritime, France, Taylor &
Hammond Colln.

DESCRIPTION. Colony unilamellar or multilamellar, each
layer about 0-27-0-41 mm thick, occasionally growing freely
with an exposed basal lamina. Overgrowths (Fig. 197) origi-
nate through intrazooecial fission from one or several closely-
spaced parental zooids; pseudoancestrula either an autozooid
(aperture 0-15 x 0-12 mm) or a kenozooid, initiating a
secondary zone of astogenetic change of increasing zooid
size. Organization fixed-walled. Zooidal apertures variably
arranged, often irregular.

Autozooids (Figs 198, 199) large, frontally elongate, about
1-6-2 x longer than wide, variable in shape; frontal wall
slightly convex, pseudopores apparently small and subcircu-
lar; boundary wall salient. Aperture of moderately large size,
a little longitudinally elongate, on average 1-1 x longer than
wide, attaining maximum width about mid-length, slightly
pointed distally; apertural rim raised, drawn to a projecting
point distally; apertural shelf narrow, tapering proximally;
hinge line short, teeth and bar not clearly visible in the
inadequately preserved material available. Operculum (Fig.
200) often preserved in-situ, convex; pseudopores not
observed due to poor surface preservation. Terminal dia-
phragms (Fig. 193) may be present just beneath apertural
shelf. Intramural buds not observed.

Kenozooids (Figs 198, 199) common, variable in number,
intercalated among other zooids in the normal budding

10 P. D. TAYLOR

ee Rd oo

199 200

Figs 197-200 Reptomultelea dixoni (Lang, 1906). 197, BMNH D7845, holotype, [Coniacian], cortestudinarium Zone, Kenley, Surrey,
England, overgrowth origin, x 23. 198, BMNH D31844, [Turonian], Dunton Green, Sevenoaks, Kent, England, autozooids, kenozooids

and an eleozooid, x 55. 199-200, BMNH D57506, Coniacian, Craie 4 M. normanniae, Etretat, Seine Maritime, France; 199, autozooids,
kenozooids and eleozooids, x 37; 200, autozooidal operculum, x 215.

sequence and also developed at anastomoses between over- frontal length: mean = 0-73 mm; SD = 0-073 mm;
growths; frontally polygonal in outline and invariably shorter CV = 10-0; range = 0-62-0-84 mm
and narrower than the autozooids. frontal width: mean = 0-41 mm; SD = 0-050 mm;
Eleozooids (Figs 198, 199, 201) moderately common, fron- CV = 12-3; range = 0:36-0:50 mm
tally elongate, 2-3 x longer than wide, longer but about the apertural length: mean = 0-24 mm; SD = 0-015 mm;
same width as the autozooids. Aperture elongate, spatulate, CV = 6:3; range = 0-21-0-26 mm
parallel-sided or slightly tapering, widest at the hinge line, apertural width: mean = 0-22 mm; SD = 0-014 mm;
rounded distally; rostral shelf extensive, depressed distally. CV = 6:2; range = 0-20-0-24 mm
pena (Fig. 201) often preserved in-situ, pseudopores not (5 zooids from BMNH D57506)
observed due to poor surface preservation. Terminal dia- ; ins
frontal length: range = 0-62-0-72 mm
phragms observed. Intramural buds not seen. cares na
Gass waknBee frontal width: range = 0-36-0-41 mm
apertural length: range = 0-23-0-26 mm
MEASUREMENTS. apertural width: range = 0-20-0-24 mm
autozooids eleozooids (5 zooids from BMNH D57506)

(10 zooids from holotype BMNH D7845)

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 71

range = 0-77-1-05 mm
range = 0-32-0-42 mm
apertural length: range = 0:38-0:59 mm
apertural width: range = 0-17-0-29 mm

kenozooids (6 zooids from BMNH D57506)
frontal length: range = 0-48-0-60 mm
frontal width: range = 0-32-0-38 mm

REMARKS. The holotype specimen lacks unequivocal eleo-
zooids, as noted by Lang (1906) in his original description of
Reptomultelea dixoni which is based entirely on this speci-
men. However, eleozooids are present in most other colonies
(e.g. Figs 198, 199, 201). Among congeneric species, R.
dixoni closely resembles R. sarissata Gregory, 1899 in the
morphology of the eleozooid which has an elongate aperture
and is often spatulate. However, although similarly pointed,
the autozooidal apertures of R. dixoni lack the distal prolon-
gations of R. sarissata apertures which give them a distinctive
ogee arch-shape (Figs 282, 283).

The proportion of kenozooids to other zooids varies greatly
between colonies of this species. In some colonies (e.g.
BMNH D31844), kenozooids occupy about the same surface
area and are more numerous than autozooids. The high
proportion of kenozooids in colonies like this one are unre-
lated to disturbances in growth caused by anastomoses of
overgrowths.

Colonies vary from lamellar to nodular in shape. Specimen
,BMNH D57505 bioimmures a perished substratum of
unknown identity which was roughly cylindrical in outline
and ornamented by tubercles.

_ frontal length:
frontal width:

DISTRIBUTION. Turonian (/ata Zone)-Coniacian (cortestudi-
| narium Zone) of southern England and northern France.

Reptomultelea filiozati (Levinsen, 1912) Figs 202-208

1912 Meliceritites filiozati Levinsen: 34, pl. 6, figs 7-10.
1990 Reptelea filiozati (Levinsen); Taylor: fig. 11.2.

MATERIAL. Lectotype (herein designated): ZMC Levinsen
Collection M13, ‘Middle Senonian’, Fécamp, Seine-
Maritime, France; the specimen shown in Levinsen’s plate 6,
figure 7. Paralectotypes: ZMC Levinsen Colln M12, M14—20,
| same details as lectotype.

Other material: BMNH D54286-7, Coniacian, Fécamp,
Voigt Colln. VH un-numbered specimens from same horizon
and locality.

DESCRIPTION. Colony unilamellar or multilamellar, each
layer about 0-3 mm thick, often becoming free with a
transversely wrinkled basal lamina. Overgrowths apparently
originate through intrazooecial fission, often from several
closely-spaced pseudoancestrulae (Fig. 206); pseudoancestru-
lae are autozooids with apertures about 0-11 mm long and
wide, sometimes operculate (Fig. 207); peri-pseudo-
ancestrular zooids number 5-6 and commence a zone of
secondary astogenetic change marked by increasing zooid
size. Organization fixed-walled. Zooidal apertures arranged
roughly in quincunx.

Autozooids (Figs 202, 205) of large size, elongate, on
average a little less than twice as long as wide, diamond-
shaped, pentagonal, rhomboidal or hexagonal in outline,
well-rounded distally; frontal wall gently convex, occupying
about two-thirds of the frontal surface; pseudopores small,
circular; boundary wall salient, clearly visible. Apertures
(Fig. 204) of moderate size, elongate, about 1-25 x longer

Fig. 201 Reptomultelea dixoni (Lang, 1906), BMNH D57505,
Coniacian, Craie 4 M. normanniae, Etretat, Seine Maritime,
France; eleozooid with in-situ operculum, x 105.

than wide, attaining maximum width about mid-length, well-
rounded distally; apertural rim slightly raised; apertural shelf
narrow, tapering proximally; hinge line with a median bar.
Operculum (Fig. 203) often preserved in-situ, slightly convex;
pseudopores not clearly visible in studied material. Dia-
phragms and intramural buds not observed.

Kenozooids (Figs 202, 205) common, especially clustered
at anastomoses between overgrowths.

Eleozooids (Figs 202, 206, 208) moderately common,
large, frontally elongate, about 2-5-3 x longer than wide,
longer but slightly narrower than the autozooids. Aperture
elongate, about twice as long as wide and twice the length of
an autozooidal aperture, more or less parallel-sided, well-
rounded distally with an extensive rostral platform depressed
beneath the level of the apertural rim; hinge line with a long
median bar. Opercula not observed in-situ. Intramural buds
not seen.

Gonozooids unknown.

MEASUREMENTS.

autozooids (10 zooids from lectotype ZMC M13)

frontal length: mean = 0-62 mm; SD = 0-049 mm;
CV = 8-0; range = 0:56-0:71 mm

Figs 202-207 Reptomultelea filiozati (Levinsen, 1912), ‘Middle Senonian’, Fécamp, Seine-Maritime, France. 202-204, ZMC Levinsen Colln
M12; 202, autozooids, eleozooids and kenozooids, x 55; 203, autozooidal operculum, x 245; 204, autozooidal aperture, x 250. 205-207,
ZMC Levinsen Colln M13, lectotype; 205, area of coalescent growth with kenozooids and autozooids, x 55; 206, four closely-spaced
pseudoancestrulae surrounded by radiating autozooids, eleozooids and kenozooids, x 55; 207, operculate pseudoancestrula surrounded by

five autozooidal buds, x 78.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 73

Fig. 208 Reptomultelea filiozati (Levinsen, 1912), ZMC Levinsen
Colln M12, Middle Senonian’, Fécamp, Seine-Maritime, France;
eleozooid, x 155.

>
4

mean = 0-34 mm; SD = 0-036 mm;
CV = 10-7; range = 0-30-0-42 mm
mean = 0-20 mm; SD = 0-009 mm;
CV = 4-8; range = 0-18-0-21 mm
mean = 0-16 mm; SD = 0-006 mm;
| CV = 3-9 range = 0-15-0-17 mm

| eleozooids (5 zooids from lectotype ZMC M13)

frontal length: range = 0-68-0-80 mm
_ frontal width: range = 0-27-0-30 mm
| apertural length: range = 0-38-0-41 mm
apertural width: range = 0-17-0-20 mm

| frontal width:
| apertural length:

/ apertural width:

REMARKS. The shape of the eleozooidal aperture is very
characteristic in Reptomultelea filiozati and enables immedi-
jate distinction from most other species of the genus. Only R.
pegma sp. nov. (see p. 85) and R. scanica sp. nov. (see p. 96)
have similarly broad eleozooidal apertures, but both species
‘are readily distinguished from R. filiozati by other means
(e.g. R. pegma has oblique autozooidal apertures with wide
japertural shelves, and R. scanica has autozooidal apertures
‘which occupy a significantly greater proportion of the frontal
surface than in R. filiozati).

According to Prof. E. Voigt, this is a rare species. It is
known to occur with certainty only in the Coniacian of the
Fécamp area of Seine-Maritime in northern France, although
the Voigt Collection in Hamburg contains a possible example
(un-numbered) from the Coniacian of Villedieu.

DISTRIBUTION. Coniacian of Seine-Maritime, France.

Figs 184, 209-216

MATERIAL. Holotype: EM RE 551.763.31.A745/1, Cenoma-
nian, Essen, Westfalia, Germany. Paratypes: VH 10426,
10548 (8 fragments), Lower Cenomanian, Miilheim/Ruhr,
Westfalia, Germany.

Reptomultelea goldfussi sp. nov.

NAME. In honour of Georg August Goldfuss (1782-1848),
the first author to describe a melicerititid species, Ceriopora
gracilis Goldfuss, 1827, later to become the type species of
Meliceritites Roemer.

DESCRIPTION. Colony (Figs 184, 211) unilamellar or multila-
mellar, each layer about 0-25-0-35 mm thick, sometimes
growing freely of underlying layers and incorporating second-
ary substrata. Ancestrula not observed. Overgrowths origi-
nate through intrazooecial fission, often from several closely-
spaced parent zooids; pseudoancestrula (Figs 212, 213) an
autozooid which may be occluded by a terminal diaphragm,
initiating a secondary zone of astogenetic change of increas-
ing zooid size. Organization fixed-walled. Zooidal apertures
arranged in regular quincunx, disrupted close to overgrowth
origins.

Autozooids (Figs 209, 212) of moderate size, frontally
elongate, about 1-5-2 x longer than wide, rhomboidal in
outline, well-rounded distally; frontal wall occupying about
half of frontal surface, slightly convex, with circular
pseudopores; boundary wall salient, variably prominent.
Aperture (Fig. 212) large, usually a little longer than wide,
attaining maximum width just proximally of mid-length;
apertural rim conspicuous but low, continuous with lateral
zooecial boundary wall; apertural shelf rather narrow (less
than 0-02 mm wide), tapering to nothing at widest point of
aperture; hinge line with median bar and low teeth at either
end. Operculum (Figs 210, 220) occasionally preserved
in-situ, surface convex; pseudopores elongated parallel to
growth direction, scattered across entire surface of opercu-
lum. Terminal diaphragms seldom observed, pseudoporous.
Intramurally budded autozooids uncommon.

Kenozooids not observed.

Eleozooids not observed, presumed absent.

Gonozooids represented by one example in the holotype
colony (Fig. 209); frontal surface slightly longer than wide,
becoming triangular in outline soon after emergence of the
distal frontal wall from the maternal aperture; margins of
distal frontal wall irregular, indented by adjacent autozooids.
Ooeciopore very large, about the same width as an autozooid
aperture but less than half the length.

MEASUREMENTS.

autozooids

(10 zooids with in-situ opercula from holotype EM RE
551.763.31.A745/1)
frontal length: mean = 0-55 mm; SD = 0-051 mm;
CV = 9-2; range = 0-48-0-63 mm
mean = 0-32 mm; SD = 0-018 mm;
CV = 5-5; range = 0-30-0-35 mm

frontal width:

P. D. TAYLOR

Figs 209-210 Reptomultelea goldfussi sp. nov., EM RE 551.763.31.A745/1, holotype, Cenomanian, Essen, Westfalia, Germany; 209,
gonozooid and autozooids, x 30; 210, autozooidal operculum, x 150.

mean = 0:26 mm; SD = 0-022 mm;
CV = 8:8; range = 0-23-0-30 mm
mean = 0-25 mm; SD = 0-018 mm;
CV = 7:2; range = 0:23-0:29 mm

apertural length:
apertural width:

gonozooids
(1 zooid from holotype EM RE 551.763.31.A745/1)

frontal length: 2:03 mm

distal frontal wall 1-80 mm
length:

frontal width: 1-80 mm

ooeciopore length: 0-11 mm

ooeciopore width: 0-24 mm

REMARKS. The large and well-rounded autozooidal aper-
tures distinguish Reptomultelea goldfussi from other species
in the genus. As noted below (p. 91), R. goldfussi resembles
R. reedi sp. nov. but is distinguished most convincingly by the
occurrence of pseudopores across the entire surface of the
operculum rather than in a crescentic row. The holotype (Figs
209, 210) differs from the paratypes (Figs 211-216) in having
somewhat taller and larger autozooidal apertures. These
differences may be astogenetic in origin because the
paratypes comprise zooids close to overgrowth originations
and probably within secondary zones of astogenetic change.

DISTRIBUTION. Lower Cenomanian of Westfalia, Germany.

Reptomultelea levinseni sp. nov. Figs 217-223

MATERIAL. Holotype: BMNH D58930 (2 fragments), Conia-
cian or basal Santonian, decipiens Chalk, Vattetot, Seine
Maritime, France, Taylor & Hammond Collection.
Paratypes: BMNH D58931 (sample), D59248, D59249, same
horizon and locality as holotype. ZMC Levinsen Colln M56,
[?Coniacian], Fécamp, Seine Maritime, France; labelled by
Levinsen ‘Clausimultelea sp.n.?’.

NAME. For G.M.R. Levinsen, author of the most important
work on melicerititid cyclostomes.

DESCRIPTION. Colony generally multilamellar, each layer

about 0-26 mm thick and occasionally growing free of under-
lying layers and with an undulose basal lamina. Ancestrula
not observed. Overgrowths originate by intrazooecial fission,
often from several clustered parental zooids depressed
beneath general level of colony surface; pseudoancestrula an
autozooid or a kenozooid budding a ring of daughter zooids,
sometimes including a gonozooid (Fig. 221), forming the
initial part of a secondary zone of astogenetic change through
which zooid size increases. Organization fixed-walled. Zooids
arranged roughly in quincunx.

Autozooids (Figs 217, 218) of large size, frontally elongate,
about 2-2-5 x longer than wide, generally hexagonal to
diamond-shaped in outline, pointed distally; frontal wall
occupying more than half of the frontal surface, slightly
convex, with circular pseudopores set in elongate depres-
sions; boundary wall well-defined, salient. Aperture (Fig.
219) of moderate size, approximately 1-5 x longer than wide,
gothic arch-shaped, pointed distally, attaining maximum
width at a point between the hinge line and mid-length;
apertural rim well-developed, pointed and raised distally;
apertural shelf narrow; hinge line short, with low teeth at
either end of a median bar. Operculum (Fig. 220) often
preserved in-situ, convex, pseudopores present but number
uncertain. terminal diaphragms may be present beneath level
of operculum. Intramural buds not observed.

Eleozooids (Figs 217, 218, 223) abundant, scattered; frontal
surface about 3-4 x longer than wide, considerably longer than
the autozooids but approximately the same width; frontal wall
occupying more than half of the frontal surface, slightly convex
and with a similar density of pseudopores to the autozooids.
Aperture elongate, 2-3-5 x longer than wide, widest a little
distally of the hinge line, with a long, narrow rostral area
becoming parallel-sided or even very slightly spatulate; rostrum
depressed distally; hinge line with a median bar proximal to
which is a depressed area; aperture width constricted proximally
by wall thickenings which join distally to form the rostral shelf.
Operculum (Fig. 223) sometimes preserved in-situ, surface
convex, especially distally. Terminal diaphragms and intramural
buds not observed.

211 212

Figs 211-216 Reptomultelea goldfussi sp. nov., VH 10426, Lower Cenomanian, Miilheim/Ruhr, Westfalia, Germany; 211, small colony
encrusting a shell fragment, x 14; 212, autozooids radiating from two closely-spaced pseudoancestrulae with apertures closed by terminal
diaphragms, x 45; 213, detail of occluded pseudoancestrulae, x 130; 214, autozooidal operculum, x 200; 215, autozooidal
aperture, x 200; 216, growing edge showing new buds with porous interior walls, x 97.

Figs 217-222 Reptomultelea levinseni sp. nov. 217, ZMC Levinsen Collection M56, [?Coniacian], Fécamp, Seine Maritime, France; autozooids
and two eleozooids with rostra resting on kenozooids, the right-hand eleozooid fouled by a small stomatoporid cyclostome; x 60. 218-222,
Coniacian or basal Santonian, decipiens Chalk, Vattetot, Seine-Maritime, France. 218-220, BMNH D58930, holotype; 218, partly
exfoliated specimen with operculate autozooids and eleozooids, <X 28; 219, autozooidal aperture, x 265; 220, autozooidal
operculum, X 265. 221, 222, BMNH D59249; 221, gonozooid originating as a first generation bud from a pseudoancestrula (lower
centre), X 82; 222, ooeciopore, x 180.

Kenozooids variable in abundance, occasionally isolated
out often aggregated, sometimes associated with eleozooids
Fig. 217).

Gonozooids (Fig. 221), known from only one colony
| BMNH D52949) which has two, longitudinally elongate,
}
wall emerging from the maternal aperture and dilating to give
subcircular inflated distal frontal wall. Ooeciopore (Fig.

MEASUREMENTS.
utozooids (10 zooids with in-situ opercula from holotype

mean = 0-66 mm; SD = 0-049 mm;
CV = 7-5; range = 0-60-0-75 mm
mean = 0:31 mm; SD = 0:019 mm;
CV = 6-2; range = 0:27-0:33 mm
mean = 0-23 mm; SD = 0-008 mm;
CV = 3-4; range = 0-23-0-24 mm
mean = 0:16 mm; SD = 0-009 mm;
CV = 5-2; range = 0-15-0-18 mm

vleozooids (8 zooids from holotype BMNH D58930)

frontal length: mean = 1:04 mm; SD = 0-114 mm;
CV = 10-9; range = 0-90-1-23 mm
mean = 0:29 mm; SD = 0-014 mm;
CV = 4-7; range = 0:27-0:32 mm
mean = 0-52 mm; SD = 0-067 mm;
CV = 12-9; range = 0-41-0-60 mm
mean = 0-18 mm; SD = 0-025 mm;
CV = 13-6; range = 0-15—0-20 mm

konozooids (2 zooids from BMNH D52949)

Frontal width:

| pertural length:

“apertural width:

frontal length: 1-70-1-80 mm
frontal width: 1-05-1-20 mm
Doeciopore length: 0-08 mm
doeciopore width: 0-11 mm

REMARKS. This new Coniacian/Santonian species is very
similar to Reptomultelea sarthacensis from the Cenomanian-
?Lower Turonian (see p. 54). The shape and size of the
autozooid and eleozooid apertures are indistinguishable in
he two species, but R. levinseni has autozooids and eleozoo-
ds with longer frontal walls, as reflected in the larger frontal
length of these zooids (mean values of 0-66 mm for autozoo-
ids and 1-04 mm for eleozooids in R. levinseni versus
J-48-0-55 mm for autozooids and 0-70—-0-81 mm for eleozoo-
ids in R. sarthacensis).

A notable feature of the only fertile colony available
(BMNH D52949) is the budding of gonozooids from
pseudoancestrulae, well within secondary zones of astoge-
netic change. In general, gonozooids in multilamellar melic-
erititids occur exclusively in zones of astogenetic repetition.
Colonies of Reptomultelea levinseni from the decipiens
Chalk occur as nodular or tube-shaped colonies, the latter
aving probably grown loosely around organic substrates. R.
sarissata and other bryozoans from here may adopt similar
rowth-forms.

DISTRIBUTION. Coniacian and/or basal Santonian, Seine
‘Maritime, France.

eptomultelea matutina sp. nov. Figs 224-229

MATERIAL. Holotype: BMNH D52565 (several fragments,
including an SEM stub, from one colony), Lower Albian

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 7a

Fig. 223 Reptomultelea levinseni sp. nov., BMNH D58930, holotype,
Coniacian or basal Santonian, decipiens Chalk, Vattetot,
Seine-Maritime, France; eleozooid with in-situ operculum, x 160.

(tardefurcata-mammillatum Zones), Shenley Limestone,
Munday’s Hill Pit, Leighton Buzzard, Bedfordshire,
England, R.J. Hogg Collection (see Owen 1972).

NAME. From matutinus, Latin for early (in the morning),
with reference to the early geological age of the species.

DESCRIPTION. Colony encrusting, multilamellar (Fig. 224),
up to 12 or more layers each about 0-20 mm thick. Ancestrula
unknown. Overgrowths develop by eruptive budding onto
colony surface from closely-spaced autozooidal pseudoances-
trulae located in depressions (Fig. 225); apertures of
pseudoancestrulae are smaller and less elongate than those of
autozooids from zones of astogenetic repetition, being about
0-15 mm long by 0-14 mm wide.

Autozooids (Fig. 226) of large size; frontally elongate, 2-3
x longer than wide, generally rhomboidal, distally extended
and well-rounded by aperture; frontal wall occupying most of
the frontal area, convex with circular pseudopores set in
elongate depressions; zooecial boundary wall prominent.
Aperture small, longitudinally elongate, about 1:2-1-3 x
longer than wide, attaining maximum width between the
hinge line and mid-length, rounded distally; apertural shelf
present; apertural rim well-developed; hinge line with

P.D. TAYLOR

Figs 224-229 Reptomultelea matutina sp. nov., BMNH D52565, holotype, L. Albian (tardefurcata-mammillatum Zones), Shenley Limestone,
Munday’s Hill Pit, Leighton Buzzard, Bedfordshire, England; 224, multilamellar colony with exfoliated layers of autozooids, x 17; 225,
depressed area with closely-spaced, operculate pseudoancestrulae, x 60; 226, autozooids growing from top left to bottom right, x 58; 227,
autozooidal operculum, 210; 228, area of coalescent growth, x 41; 229, detail showing kenozooids, x 72.

median bar, raised above general level of frontal surface so
that plane of aperture slopes slightly downwards in a distal
direction. Operculum (Fig. 227) usually preserved in-situ,
surface a little convex, pseudopores apparently absent. Intra-
mural buds and terminal diaphragms not observed.
Eleozooids absent.
_ Kenozooids (Fig. 229) mainly developed at overgrowth
origins and where growing edges from different overgrowths
anastomose (Fig. 228), occasionally present elsewhere inter-
‘spersed among autozooids.
| Gonozooids unknown.

MEASUREMENTS.
fae 2200%ds (10 zooids with in-situ opercula from holotype

BMNH DS2565)
frontal length: mean = 0-72 mm; SD = 0-055 mm;
CV = 7-6; range = 0-66—0-81 mm
mean = 0-29 mm; SD = 0-019 mm;
CV = 6-7; range = 0:26-0:32 mm
mean = 0-17 mm; SD = 0-009 mm;
CV = 5-6; range = 0-15-0-18 mm

frontal width:

apertural length:

732

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 79

apertural width: mean = 0:13 mm; SD = 0-005 mm;

CV = 3-6; range = 0:12-0:14 mm

REMARKS. This is the earliest known species of Reptomutlte-
lea, and the only species of the genus recorded from the
Lower Albian. Only one specimen is known but this is
sufficiently different from other species of the genus to
warrant recognition as a new species. The absence of eleo-
zooids is best interpreted as a primary characteristic of the
species rather than an artefact of small sample size because
eleozooids are not developed in areas of anastomosis, where
zooid crowding and disruption of the normal budding pattern
might be expected to favour their presence. R. matutina
differs from the somewhat similar R. oceani (d’Orbigny) in
lacking eleozooids and in having slightly smaller autozooidal
apertures.

DISTRIBUTION. Lower Albian of Bedfordshire, England.

Reptomultelea mitrus sp. nov. Figs 230-234

MATERIAL. Holotype: BMNH D14468, Cenomanian [prob-

233

Figs 230-233 Reptomultelea mitrus sp. nov. 230, 231, BMNH D14468, holotype, Cenomanian, Beer Head, Devon, England; 230, operculate
autozooids, xX 27; 231, detail, x 60. 232, 233, BZ 998, Cenomanian, Wilmington Sands, Grizzle Bed, White Hart Sandpit, Wilmington,
Devon; 232, autozooids, x 29; 233, three autozooidal apertures, x 80.

ably Beer Head Limestone Formation, mantelli Zone; see
Jarvis and Tocher 1987], Beer Head, Devon, England,
Bather Collection. Paratype: BZ 998, Cenomanian, Wilming-
ton Sands [a facies of the Beer Head Limestone Fm. ], Grizzle
Bed, White Hart Sandpit, Wilmington, Devon, Greenaway
Colln.

NAME. From Mitra, Latin for head-dress, with reference to
the similarity in shape of the operculum to a Bishop’s mitre.

DESCRIPTION. Colony multilamellar, each layer about 0-45
mm thick. Ancestrula unknown. Overgrowths arise through
intrazooecial fission; pseudoancestrulae are autozooids with
apertures about 0-14 x 0-12 mm in size; secondary zones of
astogenetic change marked by increasing zooid size and
elongation of apertures. Organization fixed-walled. Zooidal
apertures closely-spaced, arranged in regular quincunx, dis-
rupted close to overgrowth origins.

Autozooids (Figs 230, 231) of large size, frontally elongate,
about 2-3 x longer than wide, typically diamond-shaped in
outline but sometimes rhomboidal; frontal wall convex, occu-
pying about half of the frontal surface, with circular
pseudopores; boundary wall inconspicuous, slightly salient.
Aperture (Fig. 233) very large, elongate, about 1-5 to almost
twice as long as wide, attaining maximum width between the
hinge line and mid-length, mitre-shaped, pointed distally;
apertural rim narrow; apertural shelf narrow, tapering proxi-
mally; hinge line with median bar and low teeth at either end.
Operculum (Fig. 234) often preserved in-situ, strongly con-
vex; pseudopores elongate, numbering about 20, arranged in
an irregular crescent often confined to the proximal half of
the operculum. Terminal diaphragms may be present just
beneath level of apertural shelf. Intramural buds not
observed.

Eleozooids unknown, presumed absent.

Gonozooids not observed.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from holotype
BMNH D11468)
frontal length: mean = 0-73 mm; SD = 0-084 mm;
CV = 11-4; range = 0-63-0-86 mm
mean = 0-32 mm; SD = 0-022 mm;
CV = 7-0; range = 0:30-0:38 mm
mean = 0:35 mm; SD = 0-024 mm;
CV = 6-7; range = 0:32-0:38 mm
mean = 0-21 mm; SD = 0-017 mm;
CV = 8-1; range = 0-20-0-24 mm

frontal width:
apertural length:

apertural width:

REMARKS. Reptomultelea mitrus is characterized by the large
and high apertures which are closely-spaced. There is some
resemblance with R. bituberosa sp. nov., but the prominent
tubercles of R. bituberosa are totally absent in R. mitrus and
eleozooids have not been found in this latter species. Aper-
tures in the paratype specimen (Figs 232, 233) of R. mitrus
are less longitudinally elongate than in the holotype (Figs
230, 231), but this may be due to proximity to overgrowth
origins within secondary zones of astogenetic change. Char-
acteristic of the species are the strongly convex opercula and
typically diamond-shaped frontal outlines of the zooids. The
most prominent, proximal areas of the opercula are often
missing in the holotype following exfoliation of overlying
layers of the colony (Figs 230, 234).

Neither the holotype nor the paratype preserve their
original substrates which may have been aragonitic shells.

P. D. TAYLOR

The paratype is a ‘taco-shaped’ colony which apparently grew
over the edge of a partly enveloped tabular substratum,
possibly a bivalve shell fragment.

DISTRIBUTION. Cenomanian of Devon, England.

Reptomultelea oceani (d’Orbigny, 1850) Figs 235-242

1850 Diastopora oceani d’ Orbigny: 176.
1853 Reptelea oceani (d’Orbigny); d’Orbigny: 641, pl.
636, figs 5, 6.
non 1872 Diastopora oceani (d’Orbigny); Reuss: 110, pl.
27, figs 2, 3.
1890 Reptelea oceani (d’Orbigny); Pergens: 399.
1899 Reptelea pulchella (d’Orbigny) var. plana

(d’Orbigny); Gregory: 292 (partim), fig. 31.
non 1899b Reptelea oceani (d’Orbigny); Gregory: 295.
1906 Semimultelea irregularis d’Orbigny; Lang: 63.
21906 Reptelea oceani (d’Orbigny); Lang: 63.

MATERIAL. Holotype: MNHN d’Orbigny Collection 6561 |

(Fig. 235) (Voigt photocards 5681 and 5703), Cenomanian,

Le Havre; this specimen measures about 24 x 16 mm and is ©

an excellent match with d’Orbigny’s figure (1853: pl. 636, figs

Fig. 234 Reptomultelea mitrus sp. nov., BMNH D14468, holotype,
Cenomanian, Beer Head, Devon, England, autozooidal
operculum, x 205.

;

| SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 81

a 7a "238

239° mee cs 240

‘Figs 235-240 Reptomultelea oceani (d’Orbigny, 1850). 235, photograph of MNHN d’Orbigny Collection 6561, holotype, Cenomanian, Le
Havre, Seine Maritime, France, X 12. 236, BMNH D4387, Cenomanian, Craie chloritée, Cap de la Héve, Seine Maritime, France,

| gonozooid with associated autozooids and kenozooids, x 50. 237, BMNH D58737, Cenomanian, Cap de la Héve, autozooids, kenozooids
and an eleozooid, x 43. 238-240, VH 10447, Cenomanian, Cap de la Héve; 238, autozooids, the two in the lower right closed by terminai

| diaphragms with central depressions, x 75; 239, autozooidal operculum, x 215; 240, autozooid with broken operculum and terminal

diaphragm beneath, x 198.

5, 6), although the image is reversed in the figure.

Other material: VH 10447, Cenomanian, Cap de la Heéve,
Seine Maritime, France. VH 10446, Lower Cenomanian,
carcitanensis Zone, Carriére du Billot, Notre-Dame-de-
Fresnaye, Calvados, France, G. Breton Colln. BGS GSM
118079, 118081, 118083, 118085-7, 118103, [Lower Cenoma-
nian], Warminster Greensand, Warminster, Wiltshire,
England, purchd from W. Cunnington, 1875. BMNH D7274,
[Cenomanian, Warminster Greensand, Warminster], mentd
by Lang (1906: p. 63). BMNH D57561-6, Cenomanian,
Warminster Greensand, Warminster, J.E. Lee Colln presd
1885. BMNH D59205-6, Upper Albian, Upper Greensand,
Chert Beds (about 10 m beneath top), Beer Head, Devon,
colld by P.D. Taylor, 1985. BMNH D4387, Cenomanian,
Craie chloritée, Cap de la Héve, Seine Maritime, France, figd
as Reptelea pulchella var. plana by Gregory (1899, fig. 31).
BMNH D55059-61, Cenomanian, Le Havre, Seine Maritime,
France, S. Whiteley Colln. BMNH D58737, Cenomanian,
Cap de la Heéve, Pitt Colln. BMNH D58899, Lower Cenoma-
nian, Craie Glauconieuse, Cap de la Heve, colld by P.D.
Taylor & J. Hammond, 1985. BMNH D58924 (4 pieces),
Lower Cenomanian, Craie Glauconieuse, Bruneval, Seine
Maritime, colld by P.D. Taylor & J. Hammond, 1985.

Questionably assigned: BMNH D55565, Lower Cenoma-
nian, Glauconitic Marl, Rocken End, Isle of Wight, England,
C.W. Wright Colln, 1934.

DESCRIPTION. Colony encrusting, multilamellar (Fig. 235),
with layers about 0-3 mm thick. Overgrowths develop by
eruptive budding onto the surface of the colony;
pseudoancestrula usually an autozooid, chamber continuous
with the underlying zooid, which undergoes intrazooecial
fission to bud pseudoancestrula and encircling zooids of the
overgrowth; zone of secondary astogenetic repetition centred
on overgrowths is extensive. Ancestrula not observed. Orga-
nization fixed-walled. Autozooids commonly arranged in
approximate quincunx.

Autozooids (Figs 237, 238) of large size, frontally elongate,
on average 2:5 x longer than wide, irregularly rhomboidal or
hexagonal in outline shape, rounded to subrounded distally;
frontal wall occupying most of the frontal area, slightly convex
with circular pseudopores; zooecial boundary wall prominent
but thin. Aperture moderately large, longitudinally elongate,
1-1-1-3 x longer than wide, attaining maximum width about
mid-length, moderately rounded; apertural rim well-developed,
often pointed distally and elevated so that the plane of the
aperture slopes upwards in a distal direction; apertural shelf
slight; hinge line bowed, apparently with a median bar. Oper-
cula (Fig. 239) quite often found in-situ, convex, pseudopores
not observed. Terminal diaphragms (Fig. 238) sometimes
present, generally located well proximal to the apertural rim, in
one example underlying a broken operculum (Fig. 240),
sparsely pseudoporous and with a central depression and pore.
Intramural buds not observed.

Eleozooids (Figs 237, 241, 242) moderately abundant,
scattered among autozooids in zones of secondary astogenetic
change and repetition; frontal surface large, about 3 x longer
than wide, appreciably longer and a little wider than autozoo-
ids; frontal wall occupying about half of the frontal surface,
pseudopore density not differing from that of autozooids.
Aperture elongate, 2-2-5 x longer than wide, slightly spatu-
late or less often parallel-sided, well-rounded distally; ros-
trum forming an extensive platform. Terminal diaphragms
sometimes present (Fig. 241), occupying a variable propor-

P.D. TAYLOR

tion of the aperture, sparsely pseudoporous. Opercula not
observed. Intramurally budded autozooids may be present in
eleozooids (Fig. 242).

Kenozooids especially common in zones of secondary
astogenetic change close to overgrowth origins, in areas of
anastomosis between overgrowths, and associated with gono-
zooids (Fig. 236).

Gonozooids (Fig. 236) present in a minority of specimens,
although more than 10 examples occur in one moderately-
sized colony (VH 10446). Frontal wall equidimensional or up
to almost twice as long as wide, with a short parallel-sided —
portion emerging from the maternal aperture, inflated and
densely pseudoporous. Atrial ring present. Ooeciopore trans-
versely elliptical, twice as wide as long, a short and reflexed
ooeciostome developed in BMNH D4387.

MEASUREMENTS.

autozooids (10 zooids from VH 10447)

frontal length: mean = 0-70 mm; SD = 0-066 mm;
CV = 9-4; range = 0-62-0-83 mm
mean = 0:28 mm; SD = 0:015 mm; ~
CV = 5:3; range = 0-27-0-32 mm
mean = 0-20 mm; SD = 0-018 mm;
CV = 8-6; range = 0-18-0-23 mm
mean = 0-17 mm; SD = 0-014 mm;
CV = 8-5; range = 0-15—-0-20 mm

eleozooids (10 zooids from VH 10447)

frontal length: mean = 0-96 mm; SD = 0-081 mm;
CV = 8-4; range = 0-86-1-16 mm
mean = 0-32 mm; SD = 0-032 mm;
CV = 9-8; range = 0-30-0-41 mm
mean = 0-42 mm; SD = 0-026 mm;
CV = 6:3; range = 0:36-0:45 mm
mean = 0-18 mm; SD = 0-010 mm;
CV = 5-5; range = 0-17-0-20 mm

gonozooids (8 zooids from VH 10446)

frontal length: mean = 1:56 mm; SD = 0-219 mm;
CV = 14-0; range = 1-29-1-80 mm
mean = 1-30 mm; SD = 0-204 mm;
CV = 15-7; range = 1-07-1-53 mm
mean = 0-91 mm; SD = 0-086 mm;
CV = 9-4; range = 0-84-1-05 mm
ca 0:06 mm

ca 0-12 mm

frontal width:
apertural length:

apertural width:

frontal width:
apertural length:

apertural width:

distal frontal wall
length:
frontal width:

ooeciopore length:
ooeciopore width:

REMARKS. This species is characterized by its eleozooids
with rostra well-rounded distally and spatulate or parallel-
sided. Among other species of Reptomultelea of similar
general morphology, R. oceani can be most easily distin-
guished by the spatulate shape of most of the eleozooid
apertures.

Reptomultelea oceani is abundant in the Lower Cenoma-
nian Craie Glauconieuse of Normandy and is also common in
the approximately contemporaneous Warminster Greensand
of Wiltshire. Specimens from the Warminster Greensand
tend to have squatter autozooids than those from the Craie
Glauconieuse. A specimen from the Glauconitic Marl
(BMNH D55565) is assigned to this species with some
reservation because of the distinctly narrower eleozooid
aperture and slight differences in autozooid morphology.

DISTRIBUTION. Upper Albian-Lower Cenomanian of SW
England and Normandy.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 83

yg = : eo
igs 241-242 Reptomultelea oceani (d’Orbigny, 1850), Cenomanian,
Cap de la Héve, Seine Maritime, France. 241, VH 10447,

eleozooid with terminal diaphragm, x 115. 242, BMNH D58737,

eleozooid containing intramural autozooid, x 112.

Reptomultelea parvula sp. nov. Figs 243-248
MATERIAL. Holotype: VH 10434, Lower Cenomanian,
Mulheim/Ruhr, Westfalia, Germany, Klaumann Colln.

NAME. Parvus, small (L.), with reference to the small size of
the zooids.

DESCRIPTION. Colony multilamellar with thin layers, each
layer about 0-15 mm thick, a transversely folded basal lamina
grows free of the bioimmured substratum (?sponge) in the
holotype. Ancestrula not observed. Overgrowths (Fig. 243)
originate throug: eruptive budding onto the colony surface
(presumably as a result of intrazooecial budding);
pseudoancestrula an autozooid with a minute aperture, about
0-09 mm long by 0-06 mm wide, depressed beneath the
general level of the colony surface; autozooids in secondary
zone of astogenetic change with relatively small, distally
rounded apertures; eleozooids in secondary zone of change
with short rostra. Organization fixed-walled. Zooids arranged
in approximate quincunx.

Autozooids (Figs 245-247) small, frontally elongate, on
average slightly over twice as long as wide, typically hexago-
nal, pointed distally; frontal wall occupying over half of the
frontal surface, convex, sparsely pseudoporous; boundary
wall well-defined, salient. Aperture (Fig. 248) of small size,
on average 1-5 x longer than wide, gothic arch-shaped,
attaining maximum width somewhat proximally of mid-
length; apertural rim well-developed, prolonged into a
tubercle-like structure distally; apertural shelf narrow; hinge
bar and teeth not observed. Operculum (Fig. 244) often
preserved in-situ, medioproximal surface flat or concave,
distolateral edges slightly convex; pseudopores numbering
about 16, radially elongate, arranged in a crescent parallel to
the distolateral margins. Terminal diaphragms and intramu-
ral buds not observed.

Eleozooids (Figs 245-247) very abundant, scattered
throughout zones of astogenetic change and repetition; small,
frontal surface about 2:5 x longer than wide, similar in width
to autozooids but rather longer; frontal wall occupying about
half of the frontal surface, sparsely pseudoporous. Aperture
elongate, 2-3 x longer than wide, widest close to the hinge
line, with a long, narrow rostral area, sometimes very slightly
spatulate and rounded distally; apertural rim prolonged into a
tubercle-like structure distally. Opercula often preserved
in-situ, surface flat proximally, strongly convex in the narrow
distal part of the operculum, proximal edge thickened and
bowed; pseudopores radially elongate, arranged in a cres-
cent, becoming more widely-spaced distally. Intramural buds
not observed.

Gonozooids (Fig. 243) present in the holotype, longitudi-
nally elongate, about twice as long as wide. a variably
lengthed parallel-sided portion emerging from the maternal
aperture (Fig. 247) and becoming longitudinally ovoidal.
Ooeciopore transversely elongate, about twice as wide as
long, its proximal edge indented internally by a hemiseptum.
Atrial ring not observed.

MEASUREMENTS.

autozooids (10 zooids from holotype VH 10434)

frontal length: mean = 0-39 mm; SD = 0-033 mm;
CV = 8-5; range = 0-35—-0-45 mm
mean = 0-18 mm; SD = 0-010 mm;
CV = 5-5; range = 0-17-0-20 mm

frontal width:

Figs 243-248 Reptomultelea parvula sp. nov., VH 10434, holotype, Lower Cenomanian, Miilheim/Ruhr, Westfalia, Germany; 243, gonozo:
(left), small intrazoarial overgrowth (right), and an overgrowing onychocellid cheilostome (bottom), x 40; 244, autozooidal
operculum, x 300; 245, operculate autozooids and three eleozooids, two with in-situ opercula, x 97; 246, autozooids and an
eleozooid, x 130; 247, gonozooid origin (left), autozooids and eleozooids, x 85; 248, autozooidal aperture, x 300.

_ SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 85

| apertural length:

mean = 0-15 mm; SD = 0-010 mm;
CV = 6:9; range = 0:14-0:17 mm
mean = 0-10 mm; SD = 0-008 mm;
CV = 8-2; range = 0-09-0-11 mm

apertural width:

| eleozooids (10 zooids from holotype VH 10434)

mean = 0-45 mm; SD = 0-022 mm;
CV = 4-8; range = 0:42-0:48 mm
mean = 0-18 mm; SD = 0-021 mm;
CV = 11-5; range = 0:15-0:23 mm
mean = 0:24 mm; SD = 0-017 mm;
CV = 6:8; range = 0-23-0:27 mm
mean = 0-09 mm; SD = 0-012 mm;
CV = 12-7; range = 0-08-0-12 mm
gonozooids (3 zooids from holotype VH 10434)

frontal length: 1-34 mm (1 zooid)
distal frontal wall 0:93-1:14 mm (2 zooids)

frontal length:
frontal width:
apertural length:

apertural width:

length:
frontal width: 0-60-0-68 mm
ooeciopore length: 0-05—0-06 mm
ooeciopore width: 0-11-0-14 mm

REMARKS. This species resembles Reptomultelea sarthacensis
(dOrbigny), with which it co-occurs at Miilheim, but has
substantially smaller zooids, a difference considered sufficient to
warrant its recognition as a new species. Eleozooids seem to be
more abundant in R. parvula than R. sarthacensis, and are
present in secondary zones of astogenetic change close to
pseudoancestrulae, while autozooidal opercula have a distinct
flattened proximal area beyond which is the crescent of
pseudopores.

DISTRIBUTION. Lower Cenomanian of Mtilheim/Ruhr, West-
falia, Germany.

Reptomultelea pegma sp. nov. Figs 249-255

MATERIAL. Holotype: VH 10430, Lower Cenomanian,
Mulheim/Ruhr, Westfalia, Germany. Paratypes: VH 10431,
105436, same horizon and locality as holotype.

| NAME. Pegma, shelf (L.), with reference to the broad aper-

tural shelf.

DESCRIPTION. Colony unilamellar or locally multilamellar,
with each layer about 0-38 mm thick, typically taking the
form of a unilamellar expansion with a transversely folded
basal lamina indicating growth into free space independent of
a substratum. Colony base not observed. Overgrowths appar-
ently originate by intrazooecial fission (Fig. 250) from a
group of basal autozooids; later stages not seen. Organization
fixed-walled. Zooids arranged in approximate quincunx.
Autozooids (Figs 249, 253) of moderate size, frontally
elongate, about 1-5-2 x longer than wide, subhexagonal in
Outline with a well-rounded distal border and concave
proximo-lateral borders; frontal wall occupying much less
than half of the frontal surface, very convex with circular
pseudopores; zooecial boundary wall generally absent. Aper-
ture (Fig. 251) moderately large, a little longer than wide,
attaining maximum width about mid-length, considerably
Narrower at the proximal edge, well-rounded and strongly
depressed distally; apertural rim usually absent but some-
times developed (? signifying intramural budding); apertural
shelf very broad, attaining a width of about 0-06 mm distally,
tapering proximally; hinge line very short, raised, bowed.
Operculum (Fig. 254) often preserved in-situ, convex, with
about 20 elongate pseudopores arranged in a crescent extend-

ing from either end of the hinge line to mid-length or a little
beyond. Terminal diaphragms (Fig. 252) present in a few
autozooids slightly beneath the level of the apertural shelf,
sparsely pseudoporous. Intramurally budded autozooids not
observed, unless occasional autozooids with apertural rims
are intramural buds.

Eleozooids (Figs 249, 255) common, scattered; large, fron-
tal surface over 2 X longer than wide, widest near the distal
part of the aperture, almost twice the length and 1-5 x the
width of an autozooid; frontal wall occupying less than a third
of the frontal surface, very convex with circular pseudopores.
Aperture elongate, almost twice as long as wide, distinctly
spatulate, well-rounded distally; rostrum forming an exten-
sive platform strongly depressed distally; apertural rim
absent; hinge line raised, broken in all available specimens.
Opercula not observed in-situ. Intramural buds not seen.

Kenozooids occasionally present, sometimes associated
with eleozooids.

Gonozooids not observed.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from holotype VH

10430)

frontal length: mean = 0-48 mm; SD = 0-062 mm;

CV = 12-9; range = 0-42-0-63 mm

mean = 0-27 mm; SD = 0-015 mm;

CV = 5-6; range = 0:24-0:29 mm

apertural length: mean = 0-24 mm; SD = 0-019 mm;
: CV = 8-1; range = 0-23-0-27 mm

apertural width: mean = 0-23 mm; SD = 0-007 mm;

CV = 3-4; range = 0-21-0-24 mm

eleozooids (6 zooids from holotype VH 10430)

frontal length: mean = 0-82 mm; SD = 0-048 mm;
CV = 5-9; range = 0-75-0-89 mm
mean = 0-37 mm; SD = 0-033 mm;
CV = 8-8; range = 0-32-0-41 mm
mean = 0-55 mm; SD = 0-039 mm;
CV = 7-2; range = 0-51-0-60 mm
mean = 0:31 mm; SD = 0-029 mm;
CV = 9-5; range = 0:27-0:36 mm
[nb. apertural length is an estimated value because all avail-
able eleozooids have broken hinge lines]

frontal width:

frontal width:
apertural length:

apertural width:

REMARKS. This distinctive new species is notable for the
broad apertural shelf, sunken aperture, small frontal wall and
short hinge line of the autozooids. The apertural shelf in
particular enables a clear distinction to be made from all
other species of Reptomutltelea.

DISTRIBUTION. Lower Cenomanian of Mtilheim/Ruhr, West-
falia, Germany.
Figs 256-262

1924 Semimultelea polytaxis Voigt: 166, pl. 4, fig. 11, pl. 5,
figs 5, 6.

Reptomultelea polytaxis (Voigt, 1924)

MATERIAL. The types of this species, originally described
from Sudmerberg near Goslar and Gr. Biillten near Peine,
Westfalia, were destroyed during World War 2. As the
identity of R. polytaxis is not in doubt, it would serve no
useful purpose to designate a neotype.

Other material: BMNH D31068, D39515—7, D39524, San-
tonian, Gr. Bulten, Westfalia, Germany, Voigt Colln. VH
10304, 10453, L. Santonian, Gr. Vallstedt, nr Braunschweig,

86 P. D. TAYLOR

Figs 249-254 Reptomultelea pegma sp. nov., Lower Cenomanian, Miilheim/Ruhr, Westfalia, Germany. 249-252, VH 10430, holotype; 249,
autozooids and eleozooids, x 22; 250, autozooids showing intrazooecial fission, x 55; 251, autozooidal aperture, x 270; 252, autozooidal
aperture closed by a terminal diaphragm, x 205. 253, 254, VH 10431; 253, autozooids, x 32; 254, autozooidal operculum, Xx 270.

\
?

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 87

Fig. 255 Reptomultelea pegma sp. nov., VH 10430, Lower
| Cenomanian, Mulheim/Ruhr, Westfalia, Germany; eleozooid

with broken hinge line, x 142.

Westfalia, Germany. VH 10440, L. Santonian, Lengede-
Broistedt, nr Braunschweig, Westfalia, Germany.

DESCRIPTION. Colony multilamellar, layers variable in thick-
ness from about 0-3-0-5 mm. Basal lamina often growing free
of substratum, undulose, when weathered revealing septal
trace pattern defining long, narrow basal outlines of zooids
(about 0-65 mm long by 0-07 mm wide in VH 10453).
Overgrowth origins depressed (Fig. 257), sometimes closely-
spaced, originating by intrazooecial fission, the parent zooid
forming a central pseudoancestrula surrounded by about 6-7
buds; pseudoancestrula an autozooid or a kenozooid, com-
mencing a secondary zone of astogenetic change of increasing
zooid size. Organization fixed-walled. Zooidal apertures
irregularly arranged, especially close to overgrowth origins
and anastomoses.

Autozooids (Figs 256, 260) small, frontally elongate, about
twice as long as wide, usually subhexagonal or subrhomboidal
jin outline, well-rounded distally; frontal wall almost flat,
/pseudopores subcircular; boundary wall salient. Aperture
(Fig. 259) of small size, longitudinally elongate, on average
fapout 1:25 x longer than wide, attaining maximum width
between the hinge line and mid-length, well-rounded distally;
/apertural rim slightly raised; apertural shelf narrow, tapering

proximally; hinge line bowed with a short median ridge
between hinge teeth. Operculum (Fig. 258) often preserved

in-situ, convex; pseudopores numbering about 14, circular or
a Ittle elongated radially, arranged in a crescent. Terminal
diaphragms occasionally present (SEM-studied examples
poorly-preserved). Intramural buds not observed.

Kenozooids (Figs 256, 260) abundant, scattered among the
autozooids and eleozooids mostly in zones of astogenetic
repetition away from overgrowth origins.

Eleozooids (Fig. 260) abundant, scattered; small, frontally
elongate, generally slightly longer than the autozooids but
about the same width, tapering distally. Aperture (Fig. 261)
small, longitudinally elongate, trifoliate in outline, on aver-
age 1-8 Xx longer than wide and slightly shorter than the
autozooidal apertures; however, eleozooids with larger aper-
tures (0-29 x 0-12 mm) are present in VH 10304; apertural
rim moderately raised distally. Operculum not observed
in-situ. Intramural buds unknown.

Gonozooids (Fig. 262) represented in only one colony.
Longitudinally elliptical in frontal outline, the distal frontal
wall commencing with a short parallel-sided tube emerging
from the maternal aperture. Ooeciopore transversely elon-
gate, about 1-7 x wider than long. Atrial ring not observed.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from VH 10440)
frontal length: mean = 0-35 mm; SD = 0-045 mm;
CV = 12-9; range = 0-29-0-42 mm
mean = 0-17 mm; SD = 0-016 mm;
CV = 9-4; range = 0-15—0-20 mm
mean = 0-15 mm; SD = 0-009 mm;
CV = 6-2; range = 0-14-0-17 mm
mean = 0-12 mm; SD = 0-005 mm;
CV = 3-9; range = 0:12-0:14 mm

eleozooids (10 zooids from VH 10440)

frontal length: mean = 0-39 mm; SD = 0-042 mm;
CV = 10-8; range = 0-33-0-48 mm
mean = 0-18 mm; SD = 0-017 mm;
CV = 9-6; range = 0-15-0-21 mm
mean = 0-14 mm; SD = 0-007 mm;
CV = 5-2; range = 0-14-0-15 mm
mean = 0-08 mm; SD = 0-011 mm;
CV = 14-4; range = 0-06-0-09 mm

gonozooids (2 zooids from VH 10304)

frontal width:
apértural length:

apertural width:

frontal width:
apertural length:

apertural width:

frontal length: 1-80-1-83 mm
distal frontal wall

length: 1-52-1-61 mm
frontal width: 1-04-1-28 mm
ooeciopore length: 0-08-0-09 mm
ooeciopore width: 0-11-0-12 mm

REMARKS. Reptomultelea polytaxis can be distinguished from
the co-occurring R. canui (Voigt) by the smaller size of the
autozooidal apertures, the trifoliate eleozooidal apertures,
and greater proportion of kenozooids. Among species of
Reptomultelea with trifoliozooids, R. polytaxis differs from R.
tuberculata (d’Orbigny) in its less elongate autozooidal aper-
tures, and from R. betusora nom. nov. in the shorter frontal
length of the autozooids, longer eleozooidal apertures, and
larger gonozooids. Together with R. parvula sp. nov. (p. 82),
this has the smallest zooids among species of Reptomultelea.

Colonies of R. polytaxis may attain large size; VH 10304
has about 14 layers of zooids and measures 78 X 30 mm. This
particular colony is rather unusual in having relatively few

88 P. D. TAYLOR

Semen roe

Figs 256-261 Reptomultelea polytaxis (Voigt, 1924). 256-259, VH 10304, L. Santonian, Gr. Vallstedt, nr Braunschweig, Westfalia, Germany;
256, autozooids and kenozooids, x 75; 257, overgrowth origin, x 90; 258, autozooidal operculum, x 300; 259, autozooidal
aperture, X 300. 260, 261, VH 10440, L. Santonian, Lengede-Broistedt, nr Braunschweig; 260, autozooids, eleozooids and
kenozooids, X 55; 261, eleozooidal aperture, x 300.

-|MATERIAL. Holotype:

=~ v * v ~
A a ee eee

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 89

Fig. 262 Reptomultelea polytaxis (Voigt, 1924) , VH 10304, L.
Santonian, Gr. Vallstedt, nr Braunschweig, Westfalia, Germany;
gonozooid, x 70.

eleozooids which have longer apertures than is typical for the
species.

DISTRIBUTION. Santonian of Westfalia, Germany.

Reptomultelea pseudopalpebrosa sp. nov. Figs 263-268

VH 7057, Lower Cenomanian,
Milheim/Ruhr, Westfalia, Germany, Franke Colln.

NAME. With reference to the similarity of the eleozooids to
those of Meliceritites palpebrosa Levinsen, 1912.

DESCRIPTION. Colony multilamellar, each layer about 0-4
mm thick. Ancestrula not observed. Overgrowths originate
by intrazooecial fission, often from a group of adjacent
zooids; pseudoancestrula an autozooid, initiating a zone of
secondary astogenetic change of increasing zooid size. Orga-
nization fixed-walled. Zooid apertures arranged in approxi-
mate quincunx. Frontal walls marked by narrow, slightly
sinuous, longitudinal stripes of paler and darker hue.
Autozooids (Figs 263-264) of moderate size, frontally
elongate, almost twice as long as wide, often rhomboidal in
outline, well-rounded distally; frontal wall occupying more
than half of the frontal surface, slightly convex and with
circular pseudopores; boundary wall discontinuous, absent
along some boundaries but raised into flanges or tubercles

elsewhere, especially near the proximo-lateral corners of
apertures. Aperture (Fig. 266) moderately large, on average
1-2 x longer than wide, well-rounded distally, attaining
maximum width about mid-length; apertural rim absent
except in intramural autozooids; apertural shelf broad distally
(0-03-0-04 mm), tapering proximally; hinge line with teeth at
either end of a median bar. Operculum (Fig. 265) often
preserved in-situ, surface convex; pseudopores elongate,
arranged in an irregular crescent; sclerites deep (visible as
moulds in sediment filling zooecial chambers). Terminal
diaphragms present. Intramural autozooids common, differ-
ing from primary autozooids in having apertural rims.

Eleozooids common (Figs 263-264), distributed singly or in
groups, apparently all intramurally budded and therefore
with the same frontal dimensions as autozooids. Apertural
area comprises a flat proximal area with few or no
pseudopores, a small, D-shaped aperture broader than long
and set in an inclined plane directed proximally, and an
extensive hood-like distal area (Fig. 268) which is conspicu-
ously pseudoporous, slightly inflated and overlaps the mar-
gins of the host aperture. Opercula sometimes in-situ (Fig.
267).

Kenozooids (Figs 263-264) present.

Gonozooids known from a single example, the proximal
part of which is obscured. Distal frontal wall subtriangular in
outline, broader than long. Ooeciopore transversely elon-
gate, width more than twice the length. Atrial ring not
observed.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from holotype VH
7057)

frontal length: mean = 0-49 mm; SD = 0-040 mm;
CV = 8-1; range = 0-42-0-54 mm
mean = 0:26 mm; SD = 0-020 mm;
CV = 7-8; range = 0-23-0-29 mm
mean = (0:21 mm; SD = 0-013 mm;
CV = 6:3; range = 0-20-0-23 mm
mean = 0:17 mm; SD = 0-008 mm;
CV = 4-5; range = 0-17-0-18 mm

frontal width:
apertural length:

apertural width:

eleozooids (intramural)
apertural length:
apertural width:

ca 0:04-0:05 mm
ca 0-07 mm

gonozooids (1 zooid from holotype VH 7057)
distal frontal wall

length: 1-20 mm
frontal width: 1:56 mm
ooeciopore length: 0-09 mm
ooeciopore width: 0-23 mm

REMARKS. Although known from only a single specimen, a
variably-preserved colony 37 x 24 mm in size with at least 17
layers of zooids, this species is sufficiently distinctive to
warrant formal naming. No other species of Reptomultelea
has eleozooids with small semicircular apertures and distal,
hood-like areas of pseudoporous calcification. In this aspect,
R. pseudopalpebrosa most closely resembles Meliceritites
palpebrosa Levinsen, 1912, a vinculariiform species from the
Senonian of France. The apertural shelf is similarly broad in
the two species, but the abundant pseudopores distributed
over the entire surface of the autozooidal opercula in M.
palpebrosa differ from those of R. pseudopalpebrosa, which
are arranged in an irregular crescent. A closer affinity seems

Figs 263-268 Reptomultelea pseudopalpebrosa sp. nov., VH 7057, holotype, Lower Cenomanian, Miilheim/Ruhr, Westfalia, Germany; 26
autozooids, eleozooids and kenozooids, x 45; 264, operculate autozooids, kenozooids and eleozooids, x 70; 265, autozooidal
operculum, X 265; 266, autozooidal aperture partly occupied by sediment mould of the opercular sclerites, x 265; 267, eleozooid with
in-situ operculum, X 265; 268, eleozooid aperture and irregular distal hood, x 265.

possible between the two species than is implied by their
inclusion in separate genera. The discontinuous, broken
zooecial boundary wall visible on the colony surface of R.
pseudopalpebrosa is another noteworthy feature, as is the
occurrence of slightly sinuous longitudinal stripes of alternat-
ing lighter and darker calcification on the frontal walls. The
origin and significance of these stripes are unknown.

DISTRIBUTION. Lower Cenomanian of Milheim, Westfalia,
Germany.

Figs 269-274

MATERIAL. Holotype: VH 10429, Upper Cenomanian, St
Calais, Sarthe, France.

Reptomultelea reedi sp. nov.

NAME. In memory of Christopher G. Reed, whose prema-
ture death in 1990 curtailed his elegant studies of bryozoan
larval morphology and development.

_DESCRIPTION. Colony unilamellar, potentially multilamellar
{as indicated by presence of incipient intrazooecial budding),
layer thickness about 0-45 mm; basal lamina with closely-
spaced transverse ridges and grooves. Ancestrula not
‘observed. Intrazooecial fission present, 5 or 6 daughter buds
‘surrounding the chamber of the parental zooid (Figs
271-272). Organization fixed-walled. Zooid apertures
arranged in regular quincunx.
Autozooids (Figs 269, 270) of moderate size, frontally
elongate, 1-8-2-4 x longer than wide, outline well-rounded
distally but indented proximally by apertures of adjacent
zooids; frontal wall convex, occupying more than half of the
frontal surface, with circular pseudopores; boundary. wall
subdued salient. Aperture (Fig. 274) small to medium-sized,
“=quidimensional or a little longer than wide, well-rounded
pay. attaining maximum width between hinge line and
mid-length; apertural rim present, distally most prominent;
pertural shelf widest distally (ca 0-02 mm), tapering proxi-
mally; hinge-line bowed. Operculum (Fig. 273) often pre-
‘Served in-situ, surface relatively flat with about 16-18
elongate pseudopores arranged in a crescent parallel to the
‘disto-lateral edges. Terminal diaphragms (Fig. 271) may be
‘developed at a level beneath the apertural shelf. Intramural
utozooids may have opercula substantially smaller than
primary autozooids, closing beneath the level of the apertural
helf.
Eleozooids observed only as intramural buds within auto-
-zooids. Apertures small, longitudinally elongate, almost
‘5 X longer than wide, prominent, set in an oblique plane
nd directed proximally. Opercula observed in-situ.
Kenozooids rare.

Gonozooids unknown.

ASUREMENTS.

utozooids (10 zooids with in-situ opercula from holotype VH
0429)

rontal length: mean = 0-55 mm; SD = 0-059 mm;
CV = 10-8; range = 0-47-0-63 mm
mean = 0-26 mm; SD = 0-010 mm;
CV = 3-9; range = 0-24-0-27 mm
mean = 0-18 mm; SD = 0-005 mm;
CV = 2-6; range = 0-18-0-20 mm
mean = 0-17 mm; SD = 0-007 mm;
CV = 4-3; range = 0-17-0-18 mm

ontal width:
pertural length:

apertural width:

leozooids (intramural)

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 9]

ca 0-15 mm
ca 0-11 mm

apertural length:
apertural width:

REMARKS. Only a single specimen of this species is known, a
free, unilamellar fragment measuring 8 x 5 mm. The pres-
ence of incipient intramural buds indicates that multilamellar
growth was possible. Reptomultelea reedi most closely
resembles the Lower Cenomanian Westfalian species R.
goldfussi sp. nov. (see p. 73), especially in the well-rounded
autozooidal apertures which indent distal zooids, but R. reedi
has smaller zooids and opercula with pseudopores arranged
in a crescent and not all over the surface as in R. goldfussi. A
further difference between the two species is the presence of
intramural eleozooids in R. reedi. Well-preserved eleozooidal
apertures have not been studied, but the general shape of the
opercula suggests that the eleozooids might be of the
trifoliozooid-type.

Numerous small borings, probably circular but usually
enlarged by chipping of the edges, penetrate the upper
surface of this specimen (Fig. 269). Most occur in autozooidal
opercula, but some are in terminal diaphragms or frontal
walls. They resemble inferred predator borings described
from other melicerititid species (Taylor, 1982).

DISTRIBUTION. Upper Cenomanian of Sarthe, France.

Reptomultelea reussi (Pergens, 1890) Figs 275-280

1872 Diastopora oceani d Orbigny; Reuss: 110 (partim), pl.
- 27, fig. 3 [non fig. 2 = Reptomultelea sarthacensis
(d’Orbigny)].
1890 Semielea reussi Pergens: 399.
21892 Semielea reussi Pergens; Pergens: 278, pl. 11, fig. 7.
1899 Reptomultelea? reussi (Pergens); Gregory: 323.
1906 Reptomultelea reussi (Pergens); Lang: 63, fig. 10.

MATERIAL. Lectotype (selected herein): SMD
un-numbered, the specimen figured by Reuss (1872: pl. 27,
fig. 3) (Voigt photocard 2166), Cenomanian [plenus Zone],
Dresden-Plauen, Germany.

Other material: BMNH D36115, Cenomanian, Untere
Planer, Plauen, Dresden, Germany, Pergens Collection. VH
10428, 10433, Cenomanian, plenus Zone, Hoher Stein,
Dresden-Plauen, Germany.

DESCRIPTION. Colony unilamellar or multilamellar (Fig.
277), each layer about 0-3 mm thick, sometimes cavariiform
(Fig. 280) or with layers growing freely of the substratum.
Ancestrula, overgrowth origins and pseudoancestrulae not
observed. Organization fixed-walled. Zooidal aperture
arrangement variable, often rather irregular.

Autozooids (Figs 275, 276) of medium size, frontally
elongate, about twice as long as wide, hexagonal, pentagonal
or diamond-shaped in outline with a pointed distal end;
frontal wall occupying more than half of the frontal surface,
slightly convex, with circular pseudopores; boundary wall
salient, sometimes poorly developed. Aperture (Fig. 279) of
moderate size, about 1-2-1-3 x longer than wide, gothic
arch-shaped, pointed distally, attaining maximum width
between hinge line and mid-length; apertural rim raised,
more prominent distally; apertural shelf absent; hinge line
gently bowed, teeth and bar not observed. Operculum rarely
preserved in-situ, surface flat; pseudopores arranged in a
crescent. Intramurally budded eleozooids may occur. Dia-
phragms not seen.

Kenozooids possibly present.

92 P. D. TAYLOR

be ages
2 we

Figs 269-274 Reptomultelea reedi sp. nov., VH 10429, holotype, Upper Cenomanian, St Calais, Sarthe, France; 269, autozooids, some with
bored opercula, x 22; 270, operculate and non-operculate autozooids, x 55; 271, autozooids variously with opercula, intrazooecial fissions
and a terminal diaphragm (top left), x 55; 272, autozooidal aperture with intrazooecial fission, x 225; 273, autozooidal operculum, x 265;
274, autozooidal aperture, x 265.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

Figs 275-280 Reptomultelea reussi (Pergens, 1890). 275, 276, BMNH D36115, Cenomanian, Untere Planer, Plauen, Dresden, Germany;
275, autozooids and eleozooids, x 50; 276, autozooids with and without opercula and an eleozooid above, x 105. 277-279, VH 104
Cenomanian, plenus Zone, Hoher Stein, Dresden-Plauen, Germany; 277, abraded growing edge, x 50; 278, eleozooidal aperture, x 240;
279, autozooidal aperture, x 240. 280, VH 10433, Cenomanian, plenus Zone, Hoher Stein, cavariiform colony with worn gonozooid, x 50.

Eleozooids (Figs 275, 276) abundant, scattered or clus-
tered; small, frontal surface variable in size, usually signifi-
cantly shorter and narrower than autozooids, and almost
twice as long as wide; frontal wall occupying more than half
of frontal surface, often somewhat depressed. Aperture (Fig.
278) equidimensional to more than 1-5 x longer than wide,
variable in size, smaller than an autozooidal aperture, attain-
ing maximum width between hinge line and mid-length,
distally elevated and pointed often with concave margins
where prolonged by rostral shelf; apertural shelf lacking;
apertural rim raised, especially distally; hinge line apparently
with short median bar. Operculum rarely preserved in-situ,
flat, conspicuously depressed beneath level of apertural rim.
Intramurally budded eleozooids within host eleozooids not
observed.

Gonozooids represented by a single, poorly-preserved
example (Fig. 280). Frontally a little longer than wide, distal
dilated frontal wall rounded subtriangular in shape.
Ooeciopore destroyed.

MEASUREMENTS.

autozooids (10 zooids from VH 10428)

frontal length: mean = 0:53 mm; SD = 0-034 mm;
CV = 6-4; range = 0-48-0-60 mm
mean = 0-27 mm; SD = 0-024 mm;
CV = 8-9; range = 0-24-0-32 mm
mean = 0-20 mm; SD = 0-011 mm;
CV = 5-2; range = 0-20-0-23 mm
mean = 0:16 mm; SD = 0-010 mm;
CV = 6:3; range = 0-15—0-18 mm

eleozooids (10 zooids from VH 10428)

frontal length: mean = 0-39 mm; SD = 0-085 mm;
CV = 21-9; range = 0-30-0-57 mm
mean = 0-21 mm; SD = 0-027 mm;
CV = 12-9; range = 0-17-0-26 mm
mean = 0-13 mm; SD = 0-041 mm;
CV = 31-5; range = 0-08-0-20 mm
mean = 0-10 mm; SD = 0-017 mm;
CV = 16-7; range = 0-08-0-12 mm

gonozooids (1 zooid from VH 10433)

frontal width:
apertural length:

apertural width:

frontal width:
apertural length:

apertural width:

frontal length: ca 1-26 mm
distal frontal wall

length: ca 1-10 mm
frontal width: ca 1:05 mm

REMARKS. Pergens (1890) proposed Semielea reussi as a new
species to replace Diastopora oceani d’Orbigny sensu Reuss,
1872. His brief description is given only as a footnote to a
revision of the Cretaceous cyclostomes figured by d’Orbigny
(1851-54) in the “Terrains Crétacés’ and lacks any figures or
mention of particular specimens. There is no evidence that he
had any of his own material at hand when erecting this
species. Therefore, the original specimens described by
Reuss (1872) are taken as the syntypes of Pergens’ new
species. Unfortunately, the two figured syntypes of S. reussi
(Reuss 1872: pl. 27, figs 2 and 3) are different species: the
specimen shown in figure 2 (Voigt photocard 2814) is prob-
ably conspecific with Reptomultelea sarthacensis (d’Orbigny)
(see p. 54); that shown in figure 3 (Voigt photocard 2166) is
here chosen as the lectotype of Reptomultelea reussi (Per-
gens, 1890).

Pergens (1892) later provided a more complete description
of Semielea reussi accompanied by a figure which shows a

P. D. TAYLOR

large eleozooid unlike those present in the lectotype of S.
reussi but resembling the eleozooids of R. sarthacensis. This
figure may be of a specimen, encrusting a sponge, from the
Maastricht Museum which was on loan to Prof. E. Voigt
when I had the opportunity to examine it in November 1987.
This specimen is conspecific with R. sarthacensis. Two identi-
fied specimens of S. reussi were donated to the BMNH by
Pergens and are registered as D36115 and D36116. The first is
conspecific with the lectotype of S. reussi, whereas the second
is indeterminate. It seems likely, therefore, that Pergens’
concept of S. reussi was a broad one which included R.
sarthacensis as well as R. reussi.

Reptomultelea reussi is readily distinguished by its small
eleozooids with apertures of a similar shape to the autozooid
apertures, except that they are prolonged distally by the
presence of a short rostral shelf. Categorizing these eleozoo-
ids according to the tripartite classification of Taylor (1986a)
is difficult; they are too small to be rostrozooids, lack the
inverted T-shape of trifoliozooids, and are more pointed than |
demizooids. Perhaps they are best regarded as highly reduced _
rostrozooids. i

None of the available material of Reptomultelea reussi is |
well-preserved; all specimens are variably abraded and have
cement-obscured surfaces. Poor preservation partly explains
the high variance of the eleozooidal dimensions, but a good
deal of this variability is biological in origin.

Both the lectotype colony and specimen VH 10433 (Fig.
280) are cavariform, thereby explaining Pergens’ (1890,
1892) placement of the species in Semielea (see p. 42). Other
specimens do not have such hollow growth-forms and assign-
ment of the species to Semielea is therefore unjustified.

DISTRIBUTION. Upper Cenomanian (plenus Zone) of Dres-
den, Germany.

Reptomultelea sarissata Gregory, 1899 Figs 281-286

1899 Reptomultelea sarissata Gregory: 322, pl. 16, fig. 7.

1906 Reptomultelea sarissata Gregory; Lang: 63, fig. 9.

1912 Meliceritites sarissata (Gregory); Levinsen: 40, pl. 2,
figs 1-3.

MATERIAL. Holotype: BMNH D7106 (main specimen plus
two fragments), Upper Chalk, Beachy Head, Sussex, Gre-
gory Colln.

Other material: BMNH D8011 (main specimen plus two
fragments), D8012, [Coniacian], cortestudinarium Zone,
Seaford, Sussex, Brydone Colln. BMNH D8568, D8576,
[Coniacian], cortestudinarium Zone, Luton, Chatham, Kent,
Gamble Colln. BMNH D11020, D11224, [?Coniacian], top of
cortestudinarium Zone or base of coranguinum Zone,
Chatham, Kent, Gamble Colln. BMNH D26835-9,
D27022-4, [Coniacian], cortestudinarium Zone, Worms
Heath, Surrey. BMNH D43661-2, [Coniacian], cortestudi-
narium Zone, Seaford Head, Sussex, Rowe Colln. BMNH
D46891, Upper Chalk, Seaford, Sussex, Stuart Colln. BMNH
D59276, [Coniacian], cortestudinarium Zone, Luton,
Chatham, Kent, Rowe Colln. BMNH D58929 (7 fragments),
Coniacian/Santonian, decipiens Zone Chalk, Vattetot-sur-
mer, Seine-Maritime, France, Taylor & Hammond Colln.
ZMC M335, Chalk, ‘Chatham or Luton’, Gamble Colln (speci-
men, on a flint, figd by Levinsen, 1912: pl. 2, figs 1-3). VH
un-numbered: Coniacian, Fécamp, Seine-Maritime, France;
Coniacian, Vattetot-sur-mer, Seine Maritime, France.

» SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS

285

Figs 281-286 Reptomultelea sarissata Gregory, 1899. 281-282, BMNH D46891, Upper Chalk, Seaford, Sussex, 281, area of coalescent
growth with autozooids, kenozooids and eleozooids, x 20; 282, autozooids and eleozooid with a broken operculum, X 48. 283-285,
BMNH D7106, holotype, Upper Chalk, Beachy Head, Sussex; 283, autozooidal operculum, x 105; 284, autozooidal aperture showing
intrazooecial fission, X 95; 285, eleozooid budded from a pseudoancestrula (lower right), x 77. 286, BMNH D8568, [Coniacian],
cortestudinarium Zone, Luton, Chatham, Kent, crushed gonozooid, x 23.

DESCRIPTION. Colony unilamellar or multilamellar, each
layer about 0-35 mm thick, layers sometimes growing freely
with an exposed basal lamina. Overgrowths originate through
intrazooecial fission (Fig. 284); pseudoancestrula an auto-
zooid with aperture about 0-21 mm long by 0-17 mm wide;
peri-pseudoancestrular buds may include autozooids, eleo-
zooids (Fig. 285) or gonozooids, initiating secondary zone of
astogenetic change during which apertural size increases and
apertures become more pointed. Organization fixed-walled.
Zooidal apertures arranged in quincunx except where dis-
rupted near overgrowth origins and anastomoses.

Autozooids (Figs 281-282) very large, frontally elongate,
generally 2-3 x longer than wide, hexagonal in outline,
pointed distally; frontal wall convex, pseudopores slightly
longitudinally elongate; boundary wall salient, often incon-
spicuous. Aperture (Fig. 284) of large size, longitudinally
elongate, on average 1-6 x longer than wide, ogee arch-
shaped with a short but distinct distal prolongation with
concave sides, attaining maximum width between hinge line
and mid-length; apertural rim salient; apertural shelf narrow,
tapering proximally; hinge line with a median bar, teeth
?present. Operculum (Fig. 283) often preserved in-situ, con-
vex; pseudopores not visible, probably as a result of poor
preservation. Terminal diaphragms and intramural buds not
observed.

Kenozooids occasionally present, usually located at over-
growth anastomoses (Fig. 281).

Eleozooids (Figs 281, 285) common; frontally very large,
elongate, 2-3 x longer than wide, longer and generally a
little wider than the autozooids. Aperture elongate, widest
proximally at level of hinge line, narrowing with appearance
of rostral shelf to become parallel-sided or very slightly
spatulate, rounded and depressed distally. Opercula some-
times preserved in-situ, convex, pseudopores not visible,
probably as a result of poor preservation. Intramurally bud-
ded eleozooids observed; apertures shorter than those of host
eleozooid and less depressed distally.

Gonozooids (Fig. 286) represented by a single, broken
example budded from a pseudoancestrula. Frontally pear-
shaped, dilated frontal wall about 1-3 x longer than wide.
Proximal floor showing outlines of underlying zooids, distal
wall smooth. Ooeciopore damaged in available material.
Atrial ring present.

MEASUREMENTS.

autozooids (10 zooids from holotype BMNH D7106)

frontal length: mean = 0:92 mm; SD = 0-134 mm;
CV = 14-6; range = 0:78-1:11 mm
mean = 0-38 mm; SD = 0-038 mm;
CV = 10-2; range = 0-30-0-44 mm
mean = 0-40 mm; SD = 0-020 mm;
CV = 5-0; range = 0:36-0:44 mm
mean = 0:24 mm; SD = 0-014 mm;
CV = 5-7; range = 0-23-0:27 mm

frontal width:
apertural length:

apertural width:

eleozooids

(3 zooids from holotype BMNH D7106)

frontal length: range = 1-32-1-44 mm
frontal width: range = 0-48-0-51 mm
apertural length: range = 0-72-0-83 mm
apertural width: range = 0-32-0-36 mm

(7 zooids from BMNH D8011)
frontal length: mean = 1:16 mm; SD = 0-040 mm;
CV = 3-4; range = 1-13-1-22 mm

P.D. TAYLOR |}.

frontal width:
CV = 8-8; range = 0-41-0-51 mm
mean = 0:73 mm; SD = 0-031 mm;
CV = 4-3; range = 0-69-0-78 mm
mean = (0:33 mm; SD = 0-013 mm;
CV = 4-1; range = 0-32-0-35 mm

gonozooids (1 zooid from BMNH D8568)

apertural length:

apertural width:

frontal length: 2:13 mm
distal frontal wall

length: 1:97 mm
frontal width: 1-61 mm

ooeciopore width: ca 0-15 mm (broken)

REMARKS. Reptomultelea sarissata is a very distinctive spe-
cies readily recognized by the large, ogee arch-shaped auto-
zooidal apertures. Eleozooidal morphology suggests affinities
with R. dixoni (Lang) (see p. 69), which also has large
autozooids. R. sarissata probably has a stratigraphical range
restricted to the Coniacian (cortestudinarium Zone and
equivalents).

Like other species of Reptomultelea from chalk facies, |
colonies are generally nodular to lamellar in shape and often ||
appear to have been attached to soft-bodied organisms whose
traces may remain as poorly-defined bioimmurations on the
undersides of colonies.

DISTRIBUTION. Coniacian (?basal Santonian) of southern
England and northern France.

Reptomultelea scanica sp. nov. Figs 287-296

MATERIAL. Holotype: VH 10441, Lower Campanian, mam-
millatus Zone, Karlshamn, Scania, Sweden. Paratype: VH |
10442, 10549 (sample), same details as holotype.

NAME. After the provenance.

DESCRIPTION. Colony unilamellar or multilamellar (Fig.
290), each layer about 0-3 mm thick, usually with a trans- |
versely undulose basal lamina growing freely of the substra-
tum. Overgrowths originate by intrazooecial fission;

pseudoancestrula an autozooid, aperture about 0-10 mm |

wide; periancestrular buds commence a secondary zone of
astogenetic change of increasing zooid size. Organization
fixed-walled. apertures arranged in approximate quincunx )
away from overgrowth origins and anastomoses. Mural spines
visible in proximal parts of zooids exposed on fractured edges
of specimens.

Ancestrula, probably of this species (Fig. 288), fouling the |
basal lamina of VH 10442, short (length 0-48 mm) with large |
protoecium (width 0-38 mm), negligable distal ancestrular
tube, and aperture 0-15 mm long by 0-14 mm wide. Opercu- |

lum not preserved, although presence of a straight hinge line | |

implies that ancestrula was originally operculate.

Autozooids (Figs 289) small, frontally elongate, usually |
less than twice as long as wide and subrhomboidal in outline
shape, rounded distally; frontal wall

spicuous and mainly comprising apertural rims. Aperture |
(Fig. 291) of moderate size, occupying about half of the
frontal area, longitudinally elongate, 1-1-1-3 x longer than
wide, rounded distally, attaining maximum width between
the hinge line and mid-length; apertural rim often thickened |
at proximolateral corners of aperture; hinge line with short
median bar between low (?)teeth; apertural shelf broad

mean = 0-45 mm; SD = 0-039 mm; |}

slightly convex, | ;
pseudopores subcircular; boundary wall salient but incon- | |

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS
|

287

underside of main colony, X 88.

\distally, tapering proximally and disappearing at about the
level of maximum aperture width. Operculum (Fig. 293)
laren preserved in-situ, convex; pseudopores radially elon-
gate, scattered all over surface of operculum. Terminal
\diaphragms (Fig. 294) observed just proximal of apertural
shelf, apparently lacking pseudopores. Intramurally budded
lautozooids probably represented by zooids with smaller oper-
cula and thicker apertural rims.

| Kenozooids rare, sometimes developed immediately distal
of eleozooids.

| Eleozooids (Figs 289, 295) moderately common, scattered,
frontally elongate, about 2:5 x longer than wide, longer and
wider than the autozooids, well-rounded distally. Aperture
longitudinally elongate, generally 2-5 x longer than wide,
distally parallel-sided or slightly spatulate and rounded; ros-
trum forming an extensive platform; apertural rim raised.
Opercula not observed in-situ. Intramural buds and terminal
diaphragms unknown.

| Gonozooids known from a single example (Fig. 296).
Frontally elongate, twice as long as wide, a short parallel-
_ sided tube emerging from the maternal aperture and dilating
_ into a longitudinally ovoidal shape. Autozooids adjacent to
distal part of gonozooid are orientated parallel to margins of
_ 20nozooid. Ooeciopore transversely elliptical, twice as wide
as long. Atrial ring not observed.

| MEASUREMENTS.

a (10 zooids with in-situ opercula from holotype VH
0441)

' rontal length:

mean = 0-44 mm; SD = 0-023 mm;
CV = 5-5; range = 0:39-0:47 mm
‘ ‘rontal width: mean = 0-24 mm; SD = 0-014 mm;
a | CV = 5-9; range = 0-23-0-26 mm
' tpertural length: mean = 0-20 mm; SD = 0-006 mm;
ie) CV = 3-2; range = 0-20-0-21 mm
f asta width: mean = 0-17 mm; SD = 0-009 mm;

"| CV = 5-6; range = 0-15-0-18 mm

288

Figs 287-288 Reptomultelea scanica sp. nov., Lower Campanian, mammillatus Zone, Karlshamn, Scania, Sweden. 287, VH 10441, holotype,
_ colony fragment with newly-developed overgrowth (lower right), <x 17. 288, VH 10442, ancestrula (presumed to be of this species) fouling

eleozooids (3 zooids from holotype VH 10441 and 3 zooids
from VH 10442)
frontal length: mean = 0-76 mm; SD = 0-080 mm;
CV = 10-5; range = 0-65—-0-87 mm
mean = 0-30 mm; SD = 0-013 mm;
CV = 4-5; range = 0-29-)-32 mm

mean = 0:44 mm; SD = 0-048 mm;
CV = 10-9; range = 0-39-0-50 mm
mean = 0-18 mm; SD = 0-011 mm;
CV = 6:2; range = 0-17-0-:20 mm

gonozooids (1 zooid from VH 10442)

frontal width:
apertural length:

apertural width:

frontal length: 1-80 mm
distal frontal wall

length: 1-64 mm
frontal width: 0-90 mm
ooeciopore length: 0-09 mm
ooeciopore width: 0-18 mm

REMARKS. The youngest known species of Reptomultelea, R.
scanica is characterized by its spatulate eleozooids, high
autozooidal apertures which occupy about half of the frontal
area of the zooid, and pseudopores distributed across the
entire surface of the autozooidal opercula. Eleozooid shape is
similar to R. filiozati (Levinsen), while the distribution of
opercular pseudopores is reminiscent of R. goldfussi sp. nov.,
but other characters ensure that neither of these species could
be confused with R. scanica.

Available specimens are small fragments of larger colonies.
With the exception of one specimen which encrusts a shell
fragment (?brachiopod), original substrates are not preserved
and the basal lamina evidently grew freely into space. The
shell-encrusting colony shows the beginnings of cavariiform
growth.

DISTRIBUTION. Lower Campanian of Scania, Sweden.

98 P. D. TAYLOR

Figs 289-294 Reptomultelea scanica sp. nov., Lower Campanian, mammillatus Zone, Karlshamn, Scania, Sweden. 289-291, VH 10441,
holotype; 289, autozooids and two eleozooids, x 67; 290, growing edge of overgrowth, x 74; 291, autozooidal aperture, x 270. 292-294,
VH 10442; 292, ooeciopore, < 130; 293, autozooidal operculum, < 275; 294, autozooidal aperture with terminal diaphragm, x 270.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 99

Fig. 295 Reptomultelea scanica sp. nov., VH 10441, holotype,
Lower Campanian, mammillatus Zone, Karlshamn, Scania,
Sweden, eleozooid, x 135.

|Reptomultelea tuberculata (d’Orbigny, 1853)
Figs 297-302

: 1853 Clausimultelea tuberculata d’Orbigny: 656, pl. 784, figs
12-15.

1890 Clausimultelea tuberculata d’Orbigny; Pergens: 398.

1899 Clausimultelea tuberculata d’Orbigny; Gregory: 405.

1912 Meliceritites tuberculata (d’Orbigny); Levinsen: 35, pl.

6, figs 1-3.

MATERIAL. Holotype: MNHN d’Orbigny Collection 8203
|(Voigt photocard 3717), Senonian, Triquerville, Seine-
\Inférieure, France.

Other material: BMNH D54295, Santonian, Evreux, Eure,
'|France, Voigt Colln. VH un-numbered material: Coniacian,
Fécamp; Coniacian, Vattetot-sur-Mer; Santonian, Evreux;
|Santonian (coranguinum Zone), Aulnay-sur-Iton.

|DESCRIPTION. Colony multilamellar, each layer about 0-3

|mm thick, growing as a series of discoidal subcolonies.
. Overgrowths originate through intrazooecial fission;
pseudoancestrulae are autozooids and are followed by a
jsecondary zone of astogenetic change. Organization fixed-
. walled. Zooidal apertures variably arranged.
| Autozooids (Fig. 297) of medium size, frontally elongate,
jon average a little less than twice as long as wide, often
jhexagonal in outline with distal margin prolonged by the
japerture; boundary wall salient. Aperture (Fig. 299) of
moderate size, very tall, 1:5 x longer than wide, attaining
‘|maximum width between the hinge line and mid-length,
rounded distally; apertural rim raised; apertural shelf moder-

Fig. 296 Reptomultelea scanica sp. nov., VH 10442, Lower
Campanian, mammillatus Zone, Karlshamn, Scania, Sweden,
gonozooid, x S58.

ately wide, tapering proximally; hinge line with a median bar
and ?teeth. Operculum (Fig. 298) convex; pseudopores num-
ber about 18 arranged in a crescent. Terminal diaphragms not
observed. Intramurally budded eleozooids (Fig. 300) present
within many autozooids; aperture similar in shape but shorter
than those of primary eleozooids, a little raised distally, an
area of calcification intervening between hinge line of host
zooid and proximal edge of eleozooid aperture; operculum
not observed.

Kenozooids (Figs 297, 301) numerous, intercalated
between the other zooids and sometimes completely sur-
rounding them.

Eleozooids (Figs 297, 301) common, elongate, about twice
as long as wide, a little longer and narrower than the
autozooids, distal outline narrow where prolonged by aper-
ture. Aperture (Fig. 302) elongate, narrow, about 3 x longer
than wide, attaining maximum width at the level of the hinge
line, slightly indented laterally by rostral shelf between hinge
line and mid-length, distally pointed and slightly raised.
Opercula not observed in-situ. Intramural buds not seen
within eleozooids.

Gonozooids unknown.

MEASUREMENTS.

autozooids (10 zooids with in-situ opercula from BMNH
D54295)
frontal length: mean = 0:54 mm; SD = 0-046 mm;

CV = 8-6; range = 0-47-0-65 mm

P. D. TAYLOR

ie
ee : on
See OR, An
mf

ee “a

Figs 297-302 Reptomultelea tuberculata (d’Orbigny, 1853), BMNH D54295, Santonian, Evreux, Eure, France; 297, autozooids, eleozooids
and kenozooids, = 60; 298, autozooidal operculum, x 225; 299, autozooidal aperture, x 225; 300, two intramural eleozooids, x 140; 301,
eleozooid surrounded by operculate autozooids and kenozooids, x 105; 302, eleozooidal aperture, x 240.

SYSTEMATICS OF MELICERITITID CYCLOSTOME BRYOZOANS 101

mean = 0-29 mm; SD = 0-023 mm;
CV = 7-8; range = 0:24-0:32 mm
ipertural length: mean = 0-21 mm; SD = 0-010 mm;
| CV = 4-7; range = 0-20-0-23 mm
ipertural width: mean = 0-14 mm; SD = 0-011 mm;
CV = 7-4; range = 0-12-0-15 mm

leozooids (4 zooids from BMNH D54295)
rontal length: range = 0-54-0-65 mm
Tontal width: range = 0-26-0-30 mm
pertural length: range = 0-18-0-26 mm
apertural width: range = 0-08 mm

‘rontal width:

EMARKS. Clausimultelea tuberculata d’Orbigny, 1853 is the
‘ype species by monotypy of Clausimultelea d’Orbigny, 1853,
genus here placed in synonymy with Reptomultelea (see p.
6). The narrow, acuminate apertures of the eleozooids,
umerous kenozooids and high autozooidal apertures are
seful features in identification of the species. The eleozooids
_nvite comparison with Meliceritites gothica and similar spe-
cles; a functional, if not phylogenetic, connection seems
ossible.
The only specimen of Reptomultelea tuberculata in the
*Orbigny Collection is a large colony reaching 50 mm in
jameter and preserved on the outside of a flint. This
pecimen has been labelled ‘Type’ by E. Voigt and corre-
ponds well with d’Orbigny’s plate 784, fig. 12. Levinsen’s
1912) material of R. tuberculata could not be positively
dentified among his collection of melicerititids in the ZMC,
Ithough this collection does include a specimen from
écamp labelled ‘Clausimultelea n. sp.’. The record of R.
uberculata given by Levinsen (p. 36) from the Danian of the
aris Basin is highly doubtful as the genus Reptomultelea is
lot known to range above the Campanian.

ISTRIBUTION. Coniacian to Santonian of France.

ACKNOWLEDGEMENTS. I am especially grateful to Professor E. Voigt
Universitat Hamburg) for his unfailing encouragement, advice and
10spitality, and also for generously making his large collection freely
iwailable for study, allowing me to borrow many hundreds of
pecimens, and donating much material to the BMNH. F. K.
cKinney kindly commented on parts of the manuscript. This work
as benefited from discussions over several years with many bryozo-
logists, notably A. H. Cheetham, R. S. Boardman, P. L. Cook, and
.D. D. Bishop. F. P. Bigey and the late E. Buge were helpful during
visit to the MNHN in Paris. H. Ristedt kindly arranged loans of
pecimens from the Goldfuss Collection in Bonn. SEM work was
ssisted by A. Burgess-Faulkner, T. S. Foster and the staff of the EM
nit at the BMNH. Pat Hart printed the photographs. D. L. Dean
Smithsonian Institution) provided expert tuition in the preparation
pf thin sections.

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CONTENTS

1 Systematics of the melicerititid cyclostome bryozoans; introduction and the genera Elea,
Semielea and Reptomultelea
P.D. Taylor

Bulletin of The Natural History Museum
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The brachiopods of the Duncannon Group
(Middle-Upper Ordovician) of southeast
Ireland

MATTHEW A. PARKES

Department of Geology, University College Galway, Galway, Ireland.
(Present Address: Dept. of Geology, National Museum of Wales, Cardiff, CF1 3NP, Wales)

CONTENTS
SMODESTS coo cabadnon aantpp gS soo Rebs eaeaddacbnEBe aan OnABUbSOchace SORRERERERPCORAMGHCY poumcmmaraaanc ice ERE AEG TGEEERGE] Nespas EEG 106
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Locality information by 1” Geological Survey of Ireland sheet ...... 14 beu 19 94 Boba] Bekeassbeteson 108
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106

M. A. PARKES

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Synopsis. Brachiopod assemblages from localities within the Duncannon Group of the Leinster terrane, southeast
Ireland, are systematically described and figured. The localities were known to 19th century geologists but have
been largely ignored since then. Re-collection permits a revised correlation between the Leinster localities and
successions of adjacent terranes. The faunal assemblages are all of Caradoc age. Kildare (Grange Hill), Ballygarvan
Bridge, Greenville-Moyne, Ballykale, Carrigadaggan and Kilbride are all probably Longvillian, whilst the faunas
from Kildare Grange Hill House Cottage, Clologe, Greenville and Raheen are of probable Soudleyan, or possibly
Harnagian, age. Brachiopods dominate the Duncannon faunas. One new species, Petrocrania harperi and one new
subspecies, Leptestiina oepiki ampla are described.

The brachiopods are closely related to coeval Anglo-Welsh Province faunas, with many conspecific forms. The
origins of the genera can be found mainly in earlier migrations of Baltic Province genera, although some
Scoto-Appalachian genera are present as early immigrants, implying a mid-Ordovician phase of breakdown of
Iapetus brachiopod provinciality. The faunas occur in volcano-sedimentary sequences, reflecting their palaeogeo-
graphical position in a volcanic arc marginal to the Eastern Avalonia microcontinent. This moved northward
throughout the Ordovician, acting as a staging post for inter-provincial migrations, until its collision with Baltica and
Laurentia.

Comparison of the assemblages reveals no direct similarities with Welsh palaeocommunities, although strong
inter-locality resemblance is noted, despite the occurrence of varied lithologies. Except for the molluscan-dominated
Soudleyan Kildare fauna, which resembles that at Herbertstown, assemblages indicate a normal marine environ-
ment in moderate to deep water surrounding volcanic centres.

INTRODUCTION

This work presents the results of re-collection and
re-examination of many fossiliferous localities within the late
Llandeilo and Caradoc strata of southeast Ireland. The area
of research is confined to the NE-SW Caledonide trending
belt of volcanic and sedimentary rocks of the Duncannon
Group, extending from Co. Wicklow through Co. Wexford to
south Co. Waterford; it also includes the peripheral Kildare
inlier of Co. Kildare. The faunas are all early to middle
Caradoc in age, and they are dominated mainly by brachio-
pods, with components of trilobites, gastropods, bivalves,

bryozoans, crinoids, cystoids and orthocones.

Brachiopods are one of the most useful animal groups in
Ordovician sequences for stratigraphical and environmental
interpretation in non-graptolitic strata, so the emphasis of the
present work is on their identification and description. Trilo-
bites, although numerically a minor component of the faunas,
are also discussed, but their detailed systematics will be
presented separately by Dr A. Owen and this author. The
preservation of almost all fossil material is as internal or
external moulds and this makes identification of some ele-
ments, such as bryozoans, gastropods and bivalves, difficult.
These elements are counted and listed in the relevant faunas,

-BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

but they are not identified or figured.

Limits of research area

With one exception, the boundaries of the Leinster terrane
(Harper and Parkes 1989, Murphy et al. 1991) define the
area, within which, outcrops of the various formations which
‘make up the Duncannon Group were all examined. The
rocks of the Tramore region south of Waterford were the
‘subject of a Ph.D. research project by Hilary Carlisle at
Queen’s University, Belfast. Although an important paper
(Carlisle 1979) summarized her main work, the Ph.D. thesis
was never completed. The completion of the brachiopod
‘systematics of the Tramore Limestone Formation is in
‘progress by Carlisle, Dr D.A.T. Harper and the present
author. The only locality from which her material was lost
was Kilbride, in the Upper Tramore Volcanic Formation, and
this important locality was re-collected.

The area of research, apart from the main belt of Duncan-
non Group rocks from Arklow through Gorey and Enniscor-
thy to the Waterford region, also takes in the Wicklow-Avoca
‘volcanics and the Kildare inlier to the north-west (Fig. 1).
Although the Ordovician greywackes of the Kilcullen Group
to the west of the Leinster Granite, in west Wicklow, have
yielded some fossils in the past, after literature research and a
cursory reconnaissance, this area was not studied, since
‘Briick (1971) has thoroughly revised the known Geological
‘Survey of Ireland (G.S.I.) fossil localities.

Outside this area in the Iapetus suture zone of eastern
Ireland, the present research has involved some restricted
fieldwork and literature work to draw comparisons and
contrasts between the faunas of the Leinster terrane and
those of the Bellewstown and Grangegeeth terranes. The
only case where detailed investigation has taken place
involves a new record of shelly fossils in the Llanvirn Hill-
town Formation of the Bellewstown terrane (Harper et al.
1991).

Revision of Duncannon Group faunas

In the Caledonides of western Europe and the eastern United
States and Canada, many areas have been the subject of
detailed modern palaeontological studies, often resulting in
monographic treatment of the more important fauna. In the
case of Wales, the Welsh Borderland and Girvan in Scotland,
many Ordovician successions have been revised. These
include the Bala area of North Wales (Williams 1963), the
Shelve district of Shropshire (Williams 1974), Girvan in
southwest Scotland (Williams 1962), Anglesey (Bates 1968),
mid and southwest Wales (Lockley & Williams 1981, Will-
iams ef al. 1981), various areas of North Wales (Pickerill &
Brenchley 1979, Lockley 1980, Hiller 1980 and Bates 1969)
and the type upper Caradoc of Shropshire (Hurst 1979a).

Although some localities in Ireland have received modern
‘axonomic treatment, such as the Tourmakeady Limestone in
Co. Mayo (Williams & Curry 1985) and the Portrane Lime-
stone in north Co. Dublin (Wright 1963, 1964), the area
-onsidered here has received only partial revision and atten-
fo Brenchley et al. (1977) completed a reappraisal of
several Caradoc localities in eastern Ireland, including Sliev-
roe near Rathdrum, some sites around Enniscorthy (Green-
lle, Greenville-Moyne) and the successions at Bellewstown
ind Grangegeeth; their paper was a fundamental resource for
he present work.

107

The present work repairs an omission in providing a
modern description of the faunas in the southeast of Ireland,
an area very poorly known by comparison with coeval
successions in other parts of Ireland and Britain. In terms of
biogeographical models the Leinster terrane occupies a piy-
otal position in cross Iapetus migrations, being the most
external or marginal area of the Eastern Avalonia microcon-
tinent and occupying a progessively more axial position
within the closing Iapetus ocean in the Ordovician. As a chain
of volcanic islands the Duncannon Group environments
provided staging posts in the dispersal of shelly benthos with
larval juvenile stages. The localities described herein are
important in charting the migration of different species
between the platform provinces of Laurentia, Baltica, Gond-
wana and the microcontinental terranes including Avalonia.

Although, with few exceptions, the existence of the faunas
described here was known to geologists in the 19th century,
progress in understanding palaeontological concepts and the
concomitant increase in differentiation of species has been
such that the faunal lists published by the early collectors are
now of little more use than as a provisional guide. One
‘species’ of the 19th century may now be recognized as
comprising three or four different genera. Examples include
‘Leptaena sericea’ for plectambonitoid genera, ‘Orthis calli-
gramma’ for impunctate orthoid genera and ‘Orthis testudi-
naria for punctate orthoids.

HISTORY OF RESEARCH

19th century research

The major reference to the faunas is that of M’Coy (1846),
whose description of the fossils collected by many workers
under the direction of Sir Richard Griffith (in his attempts to
make the first geological map of Ireland) was done largely
without knowledge of the localities or lithologies and was
thus a considerable achievement. The efforts of Griffith, and
his relationship with the official Ordnance Survey, as well as
to the geological community in Ireland, is a fascinating story
related by Herries Davies (1983), who has made clear that in
many aspects the lead in geological mapping and thinking
came from the G.S.I., and this affected progress in the
Geological Surveys of England, Scotland and Wales.

In the 19th century some major works describing Irish
geology and palaeontology included the third edition of
Siluria (Murchison 1859) and Davidson’s fine monographs of
British ‘Silurian’ Brachiopoda (1853, 1866, 1867, 1869, 1871,
1883). In addition, noteworthy works include those of Rey-
nolds & Gardiner on several specific areas including the
Kildare Inlier (1896). Also important was Reed, who pub-
lished papers on the Tramore area of Co. Waterford (1895,
1899, 1900).

1900-1950

After the initial mapping of Ireland was completed by the
G.S.1. in 1890 with the publication of 1” Sheet 10, there was
very little new research of note or new interpretations in the
following 70 years. In 1939 the Geologist’s Association pub-
lished a collection of papers on S.E. Ireland, including
Hallissy (in Smyth, 1939) on the present study area. A

108

significant precursor to new investigations was a review paper
by J.C. Harper (1948).

1950-1992

After the middle of the 20th century there were 3 main
‘schools’ of research developed in relation to the geology of
S.E. Ireland that were important to this research.

J.C. Harper, based in Liverpool. Stemming from the
interest and research of J.C. Harper in Irish Lower Palaeo-
zoic geology, a series of papers by him and colleagues was
presented. The most significant for this study is Brenchley er
al. (1977), revising some successions in eastern Ireland based
on new collections of fossils from Greenville, Enniscorthy,
Slieveroe and Grangegeeth. Others of note are Harper &
Rast (1964) on the Bellewstown succession; Crimes & Cross-
ley (1968) and Brenchley & Treagus (1970) on the Courtown
succession; Brenchley et al. (1967b) on the Tagoat faunas in
the Rosslare terrane; Harper (1952) and Brenchley ef al.
(1967a) on the Grangegeeth inlier and Romano (1980a,
1980b) on the eastern Ireland Ordovician inliers. France
(1967) also listed Caradoc fossils from Balbriggan.

Queen’s University Belfast. A succession of work on
Ordovician rocks in Leinster was also completed under the
guidance of Alwyn Williams in Queen’s University Belfast.
Aside from Wright’s researches at Kildare there was work by
Carlisle (1979) on the Tramore area, Hiller (1971) on the
Courtown rocks, and Mitchell (Mitchell et a/. 1972) on both
areas. Mitchell (1977) also subsequently completed a major
revision of the Pomeroy inlier in Co. Tyrone.

Geological Survey of Ireland. Under G.S.1I. instigation the
Leinster area was remapped. Gardiner (1967) remapped the
Duncannon area of S. Wexford and elucidated the structure
(1970) and stratigraphy (1974) of the region. The area to the
north but not adjoining, between Wexford and New Ross,
was remapped by Shannon, who described the stratigraphy
and sedimentology (1978, 1980), structure (1977, 1979b) and
petrology (1979a). More recently the area to the north in Co.
Wexford has been remapped by Geraghty (1989) and the area
further north, extending to Co. Wicklow, was the subject of
an M.Sc. thesis by Martinez (1987). This Wexford research
and work by Downes (1974), Boland (1983) and Carlisle
(1979) in the Tramore region have recently been compiled by
the G.S.I. and new maps are in production. Also, Briick ef al.
(1978, 1979) refer to many publications based on aspects
other than palaeontological research.

Recent research. This has been focussed on terrane tecton-
ics and its application to understanding the development of
the Irish Caledonides within the Iapetus Ocean (Murphy et
al. 1991). Harper & Parkes (1989) have outlined the palaeon-
tological constraints on the definition and development of
Irish Caledonide terranes.

INFORMATION SOURCES FOR FOSSIL
LOCALITIES

The primary source of information for tracing fossil localities
in this study were the memoirs of the G.S.I., which list all
fossiliferous sites encountered during the G.S.I. mapping
programme in the mid 19th century. Locality information is
restricted to a townland name (a townland is a small land

M. A. PARKES

area) and a quarter sheet of the six inch (1:10560) series of
topographic Ordnance Survey maps. However, some locali- |
ties are misplaced.

Another primary source of data was M’Coy’s (1846) Syn-
opsis of the Silurian Fossils of Ireland. The locality informa-
tion given by M’Coy was vague, with a townland name being ©
the smallest area unit used to identify sites. This may explain
why different faunas and lithologies were lumped together
under one townland name. One of the main workers who
collected for Griffith was John Kelly, who recognized that the |
locality information given by M’Coy was poor and gave more
precise details of known localities (Kelly 1860).

Apart from these major sources, published works detailed
in relevant sections provided further information, the most |
important of these being Brenchley er al. (1977). Significant
information was obtained by direct contacts with other
researchers and local farmers. The thoroughness of the
G.S.I. mapping programme and of the other 19th century
geologists is clearly shown by the fact that no new fossil _
localities were found despite extensive fieldwork.

LOCALITY INFORMATION BY 1”
GEOLOGICAL SURVEY OF IRELAND SHEET

Figure 1A shows the 1” geological sheets examined com-
pletely or in part in this research. This section gives all
relevant information on the fossil localities on each sheet,
whether or not any collection was made by the author, and
the current status of the site where known. Fig. 1B is a
location map of the main localities collected, and the sections
illustrated on Fig. 14 (p.126) plus other locations discussed in
the text.

Sheet 119

The only Ordovician fossil localities on this sheet are dis-
cussed by Baily (in Jukes et al., 1858). They all occur within
the Kildare Inlier and have been revised (Parkes and Palmer
1994).

Sheet 120

Seven localities are listed in the memoir for this sheet (Hull
1880). They were all examined recently by Brtick (1971) and
dealt with thoroughly. No further work was attempted for
this study, and no new localities are known.

Sheets 121 and 130

This memoir (Jukes & Du Noyer 1869) does not tabulate
fossil localities. Four ‘Lower Silurian’ localities are referred
to, all in the area of Rathdrum, Co. Wicklow.

(i) Rathdrum Hill, Kilecommon, Co. Wicklow, 30/1 or 3/30
(of the 6” series) in grey sandy beds. This locality was not
found on G.S.I. 6” fieldsheets and no fossiliferous rocks
were located in the area in reconnaissance fieldwork.

(ii) Quarry near Rathdrum Bridge, on road to Glenealy and
Wicklow, townland of Glasnarget north, Co. Wicklow 30
(?) in cleaved slate. The locality details are somewhat
vague and the site was not found on the G.S.I. 6°
fieldsheets, but is believed to be a quarry now occupied

“-BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

=
14014 N44 F
0a rey

ew
aA Grangegeeth Bort
48 | 40 50 Zz Drogheda eae
e
"69 Ge Hans Bellewstown, ‘\ BALBRIGGAN
81 | (@ a WA
ro a9 | 90 | : a1 | 92 | os LAMBAY
| 101 102 | -— ISLAND
5 a O >
ino | at | oo PORTRANE
22) Fe
Uy, dy]
[oui
Rp

167 | 168 [isavti7o

fee | 181

KILDARE
INLIER

Leinster
Granite

e Portlaoise

(Wicklow

RATHDRUMe

Leinster
Granite

Gorey
Q URTOW
BALLYKALE ° COORTIONN

Kilkenny

CLOLOGE,

GREENVILLE SS
e

Enniscorthy® y
RAHEEN,

Clonmel
e

=
glare
Te oF ecane

* Dungarvan

he 1 A, index map of Geological Survey of Ireland 1-inch sheets examined in whole or part for revision of fossil localities. B, location map
for the main sections shown on Fig. 14 (p. 126), and other localities in the Leinster terrane and adjoining terranes.

109

110

Table 1

and 149, with present status.

Localities from G.S.I. Memoir examined in Sheets 148

Loc. ‘'/4 sheet County & Situation & geological

no. of6’map townland formation & 1” map
Wexford Sheet 148

1 16/1 Clologe, Upper From debris, in field close
to road, from Norris
Mount to Milltown, Imile
S of Camolin; light grey &
brown slates, & tuffose
rock. See later section.

2 20/1 Killabeg Quarry on bank of R.
Bann, | mile S of Clone
Wood & 3 miles S of
Ferns; black slates.
Graptolite loc. — not
examined.

3) 20/2 Ballydonegan On road from The Harrow
to Tinnacross, 1 mile SW
of The Harrow; grey
shales. No exposure found
here.

Wexford Sheet 149

4 7/4 Kildermot See Courtown localities.

5 7/4 é :

6 7/4 me 4

7 7/4 Ballymoney, Lr.

8 7/4 Seafield ‘

9 11/2 Ballykale See later section.

10 11/2 Coolnaveagh About 2 miles S of Gorey a
little W of Ballinatray
Bridge; black slates. Not
located.

11 11/3 Ballydaniel One mile W of
Balloughter; bluish grey
compact altered rock. No
fossils found here.

12 11/4 Clogh and A little SE and SW of
Clogh.

Frankfort See later section.

13 12/1 Ballinatray, Lr. See Courtown localities.

14 12/1 i sf

15 12/1 ih M2

16 12/1 Seamount ‘

17 12/2 Duffcarrick "4

18 12/3 Coolnahinch A little SW of Ballywalter
House, 2-5 miles S of
Gorey. Not traced.

19 12/4 Seamount Graptolite locality in

Ribband Group — not
examined.

by a religious grotto about 100m from the bridge at
Rathdrum. However, no fauna was found in the very
slaty rock there.

(ii) Wicklow 30/3 one mile on road from Rathdrum to
Redcross. This was considered too obscure and the
supposed specimens too poor even in 1869 and no
attempt was made to trace the locality in this study.

(iv) An old road cutting in the townland of Slieveroe, Co.
Wicklow 30/1. This is discussed in brief below.

M. A. PARKES

Table 2 Localities from the G.S.I. Memoir examined in Sheet 158,
with present status.

Situation & geological
formation & 1” map

Loc. 1/4 sheet
no. of 6” map

County &
townland

Wexford Sheet 158

1 19/4 Kiltrea Graptolite loc. See
Brenchley et al. (1967).
Quarry near road, a little S
of Moyne House, 1-5 miles
N of Enniscorthy; dark
grey slates. Quarry now
infilled — no exposure at
all. See Brenchley et al.
(1977) for faunal lists from
collecting in that revision.
A little NE of Moyne
House, 2 miles N of
Enniscorthy; light bluish
shales, weathering brown.
No exposure at present.
Old quarry, 1 mile N of
Enniscorthy; dark grey
shales. Fauna collected
here in large blocks from
newly ploughed strawberry
field immediately adjacent
to quarry. See later
section.

See later section.

Between Ballynabarny
House

and White’s Bridge, 1-5
miles NE of Enniscorthy;
dark grey slates. This site
was not examined in this
study or by Brenchley et
al. (1977).

A little N of Ballybrennan
House, 1-5 miles W of

2 20/3 Moyne Upper

3 20/1 Moyne Lower

4 20/3 Greenville &

Moyne Upper

5) 20/3
6 20/3

Greenville
Clonhasten &

Ballynacarny
boundary

7 31/2 Ballybrennan

Clonmore, & about 6 miles |

|
|
|

SW of Enniscorthy; grey
shales and grits. Shannon
(1979a) recorded fossils
here but no exposure is
now available.

Near Chapel Village, 6
miles SW of Enniscorthy;

So ivi |
light grey shales & grits. |

Raheen

See later section.

Sheet 129

The memoir for this sheet (Mitchell 1884) has one palaeonto-
logical note by W.H. Baily concerning the only known
locality at Ballintaggart, Co. Kildare. This was examined by
Briick (1971) in his examination of fossil localities west of the
Leinster granite, and is not included here.

Sheets 138 and 139

Hull (1888), author of the G.S.I. memoir, mentions the only
fossil locality in the text (p.8). A graptolite locality with
poorly preserved forms ‘allied to, or identical with Grap-
tolithus Sedgwickii’ was known a short distance NW of
Arklow. The author has recently traced and curated these

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND
| Table 3 Localties from the G.S.I. Memoir examined in Sheet 168,

with present status.

County &
townland

Situation & geological
formation & 1” map

Loc. '/4 sheet
no. of 6” map

Waterford Sheet 168

111

Table 4 Localities from the G.S.I. Memoir examined in Sheet 169,
with present status.

Loc. ‘'/4 sheet
no. of 6” map

County &
townland

Situation & geological
formation & 1” map

Wexford Sheet 169

} 1 9/4 Gibbet Hill New road cutting on S side
of R. Suir, a little NW of
Waterford; black
argillaceous slates.
Graptolite locality, not
examined.

Old quarry at the back of
Farm House, 3 miles S of
Waterford, on the road to
Clohernagh Bridge; brown
calcareous impure

. limestone. G.S.I. 6”
fieldsheets were not
accessed for this area and
it was not traced.

Rocks on shore, a little N
of Newtown Head,
Waterford Harbour; dark
grey concretionary shales.

| Owen et al. (1986) have
revised the trilobite fauna.
D.A.T. Harper is revising
the brachiopod assemblage
from here.

1217/4 Killnre

Raheen

specimens in the G.S.I. collections. They have been identi-
fied by Dr. A. Rushton, and indicate an early Ordovician,
possibly Arenig age (John Morris G.S.I., pers. comm., 1994).

‘Sheets 148 and 149

This memoir (Hardman 1887) has palaeontological notes by
W.H. Baily, who tabulated the localities. These are repro-
‘duced here with relevant information from this revision
‘(Table 1).

| Sheets 158 and 159

W.H. Baily contributed palaeontological information io this
‘memoir (Kinahan 1882), again tabulating the fossil localities
‘and listing the fossils collected. These are listed here with
applicable information from this work (Table 2). All are from
|Sheet 158; none were known from Sheet 159.

Sheets 167, 168, 178 and 179

This memoir (Du Noyer 1865) also has palaeontological notes
by W.H. Baily, who listed the fossils collected and tabulated
the localities on Sheets 168, 178 and 179 (none were known
\from Sheet 167). The area of Sheet 167 is largely Caradoc or
Silurian and has been studied by Penney (1980), who dis-
‘cussed recent faunal dating (p.319), including a Caradoc
|graptolite record from the Ross Formation. Sheet 178 to the
west of the main Tramore volcanics is also outside the scope
of this research as discussed in the Introduction. Similarly,
Sheet 179 is all outside the confines of the present project and
the subject of revision by D.A.T. Harper, H. Carlisle and

Oo
n
2
=
&
iS)

About 1-5 miles SE of
Ballynabola; grey shales.
See later section.

A little N of Newbawn, 1-5
miles SE of preceding
locality; grey shales. No
fossiliferous exposure
found here. A new quarry
exposure with fossils was
recently reported to me
(pers. comm. M. Allen)
but not visited in this
research.

A little S of Ballygarvan
Bridge, 3 miles SW of
preceding locality; grey
shales. See later section.
Rocks on shore a little W
of Ballymadder Point; dark
grey shales. Reconnaisance
visit only made here. No
fossils located. Dan
Tietzsch-Tyler (pers.
comm. 1986) did not find
significant fossils while
mapping the area in detail.
Rocks on shore W of
preceding locality; grey
micaceous and argillaceous
shales. Same comment as
loc. 4 applies.

Carrigadaggan

Newbawn

3} 40/1 Ballygarvan

4 45/4 Ballymadder

5 45/4 Loftusacre

myself. The area is described by Carlisle (1979).

However, the Kilbride locality discussed below is within
the confines of Sheet 179 and adjacent to localities 13-16. It is
worth noting the confusion caused in the past by the similarity
of the names of Newtown Head in Waterford Harbour
(locality 3 — Raheen, but not the Raheen near Enniscorthy)
and that of Newtown Cove, Great Newtown Head and
Newtown Glen, all on the west side of Tramore Bay, fossilif-
erous strata being found at all locations. Sheet 168 contains
three localities, shown in Table 3.

Sheets 169, 170, 180 and 181

Kinahan (1879) wrote the memoir to the four sheets covering
southeast Co. Wexford, with W.H. Baily again contributing
palaeontological information in tabulated form. Sheet 169 is
the one relevant to this study. The table of ‘Lower Silurian’
localities is reproduced here with updated information (Table
4).

Sheet 170 has two listed localities which are part of the
Rosslare terrane and whose faunas were described by
Brenchley et al. (1967b). These are currently being reassessed
by Harper & Bates (in prep). Sheet 180 has only one
graptolite locality, visited but not yielding any specimens;
recent workers have not found any trace of them and suggest
the deformation is too strong to preserve fossils (Gardiner

112

1967: 6). Sheet 181 has no fossil localities at all.

A further significant point to note in connection with the
composite list of fossils collected from these sheets is that in
the sections on ‘Lower Silurian’ trilobites, brachiopods and
graptolites many species are recorded from Locality 12. This
is definitely a Carboniferous locality. According to the G.S.1.
Map Curator, A.G. Sleeman (personal communication,
1988), these records are actually from Localities 16 and 17,
which are both in the Tagoat area of the Rosslare terrane. He
detailed many further complexities resulting from ‘some
rather sloppy curating going on in the 19th century’. These
errors clearly show the need for caution in utilizing the
existing faunal lists alone in modern interpretative work.

DETAILED LOCALITY INFORMATION

The Kildare Inlier, Co. Kildare (1:126720 —
SHEET 16, N724175 — Horizons 1 & 2, N724179
— Grange Hill Cottage)

This inlier is described in detail elsewhere (Parkes and
Palmer 1994). Only two main horizons and one minor one
have been sampled extensively for the present work. All are
of Caradoc age and on the flanks of the andesites of Grange
Hill. The three localities sampled are shown in Fig. 2. The
older fauna at the back of the ruined farm cottage, called
Grange Hill House Cottage herein, is on the northern side of
Grange Hill, on the edge of the common land. Grange Hill
Horizons 1 and 2 are located only a few metres apart on the
lowest slopes of Grange Hill in and just above an old obsolete
field boundary.

The oldest reference to the inlier seems to be M’Coy
(1846), who listed many of his species from ‘the Chair of
Kildare’ (in the townland of Carrickanearla). This has led to
much subsequent confusion since the townland includes rocks
of Caradoc and Ashgill age and fossils from different horizons
were treated together.

In 1858 the G.S.I. memoir to Sheet 119 (35 NE) was
published (Jukes et al. 1858). A separate list was given for
fossils found at Grange Hill House Cottage, but the identifi-
cations were not indicative of significant differences between
that locality and the combination list for the Kildare Lime-
stone and Grange Hill. These fossils have never been
described although Williams et al. (1972) stated it was a
Soudleyan fauna. Wright (1970) published a study of the
inarticulate brachiopod Orthisocrania divaricata, which is
found only in the Caradoc siltstones dated as Longvillian on
the basis of faunal similarity with the Gelli-grin Group of
Bala in North Wales. Wright’s list is the only modern
reference to the Grange Hill (Horizons 1 & 2) fauna, which is
described fully herein.

Kilbride and adjacent localities, Co. Waterford
(1:126720 — SHEET 23, S578050)

The Kilbride locality is not a G.S.I. nor a Griffith locality but
is one of the few recently discovered fossil sites in the area. It
was found and collected first by Hilary Carlisle in the course
of her doctoral research. Carlisle (1979) described the stratig-
raphy of the Tramore area, Co. Waterford, and listed the
genera from Kilbride. Although Carlisle’s work was never

M. A. PARKES

completed, the material she collected was largely saved by Dr
D.A.T. Harper with the intention that they should jointly
complete the taxonomic study of the Tramore faunas.
Although the Tramore Limestone Formation collection is
currently under study by Harper, Carlisle and myself, the
Kilbride material was lost. Extensive re-collection of this
stratigraphically significant locality in the Upper Tramore
Volcanic Formation was a high priority in the present study.
The locality is shown in Fig. 3 and a detailed survey of the

quarry is shown in Fig. 4, which pinpoints the position from —

which the re-collection was made.

In Du Noyer (1865:18) four localities are listed by Baily
which are adjacent to Kilbride on 1” Sheet 179. These are
Towergare (13 & 16), Munmahoge and Lisduggan (14) and
Munmahoge (15), all townlands half a mile to the north of
Kilbride. For all four localities the memoir descriptions are
vague. Recourse to the 6” G.S.I. fieldsheets was necessary
since the localities are not even indicated on the 1” Sheet 179.
The original fieldsheets have been replaced by photographic
copies in the G.S.I. and only limited information could be

obtained. Munmahoge (15) was examined closely but the |

other localities were not traced. No fossiliferous exposures
were found. However, the faunal lists are short with only
three or four species other than the ubiquitous bryozoan
‘Stenopora fibrosa’, although at Munmahoge (15) abundant
specimens are indicated.

Ballykale, Co. Wexford (1:126720 —- SHEET 19,
1147570)

This locality was recorded as a G.S.I. locality (No. 9) in the

Sheet 149 memoir (Hardman 1887). It was noted as being |
‘one mile and a half south of Gorey; tuffose rock’. The |

quarter Sheet 11/2 of the G.S.I. fieldsheets showed the
locality, although it is possible to confuse it with any of
several adjacent localities such as Coolnaveagh (No. 10),
Coolnahinch (No.18), or a number of other sites indicated by

asterisks on the 1” Sheet 149, which are not easily correlated —

with the memoir table.

Kelly (1860) noted the locality as being ‘2 miles south of
Gorey, on the east side of the road. The locality is nearly
surrounded by a felspathic protrusion of yellow rock, such as
is frequent thereabouts’. The actual position of the collection
made by me is shown in Fig. 5. The locality has received no
attention since the G.S.I. memoir. Field investigation did not
reveal any exposure, but one large block in the base of the
wall bounding the farm road was found to be packed with
fossils, a nearly monospecific assemblage of Bimuria cf.
dyfiensis Lockley (p.155). Although not in situ it is believed
to be of local origin. The rock is a very tuffaceous mudstone
and has been relatively strongly deformed, but the flattened
fossil moulds are clearly identifiable.

Clologe Upper, Co. Wexford (1:126720 - SHEET
19, TO51509)

As locality 1 in the memoir for Sheet 148 (Hardman 1887),
this site was also identified clearly on the G.S.I. 6" field-
sheets, on quarter Sheet 16/1 of Co. Wexford. The memoir
records that fossils came ‘from debris, in a field close to the
road from Norris Mount to Milltown, one mile south of
Camolin; light gray and brown slates, and tuffose rock’.
Re-investigation of the area, shown on Fig. 6, failed to locate

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

Etrrri2=t=;

gaa cee fossil

GRANGE HILL

4
743

Or Be

fossil localit
. y Grange Hill

Horizont

Chair of Kildare ,

Wize
Earl's Wel! on

track

- electricity ~ a rough 40°” contour
transmission ~ ~/ grazing - line
cable

| fo)

| river or 2" buildings
| road or FY stream ve boundary [P,4/woodtand ,

Fig. 2 Locality map of the three fossil localities in the Kildare inlier; Grange Hill House Cottage, Grange Hill Horizon | and Horizon 2.

113

114

M. A. PARKES

Fig. 3 Locality map of the Kilbride locality. Key as in Fig. 2.

any in situ exposure. However, collecting amongst loose
blocks in the steep wooded slope within a restricted area
yielded a large fauna comparable with that listed by Baily (in
Hardman 1887). The fossils were found most frequently in
blocks of tuff but others came from slaty siltstones and
mudstones. The preservation is generally poor and identifica-
tion proved difficult beyond generic level. The similarity of
the fauna and lithology to that described in the memoir, and
the limited section of slope where fossiliferous blocks were
found, suggest that they are from the outcrop below the
surface drift.

Carrigadaggan, Co. Wexford (1:126720 - SHEET
23, $313240)

Despite being one of the most fossiliferous localities in the
Duncannon Group this site has received scant attention in the
last 100 years. Its exact position is shown on Fig. 7. M’Coy
(1846) gives this locality for many of his species — an
impressive list of 16, mainly brachiopods and trilobites, but
including the rhombiferan cystoid Echinosphaerites granula-
tus aS an ‘extremely common’ element of the fauna. The
faunal list given in the G.S.I. memoir (Kinahan 1879) is even
more comprehensive. Thirty-eight genera are listed, again
mainly brachiopods and trilobites but also several bivalve and
gastropod species, as well as bryozoans, orthocones, conu-

lariids and Echinosphaerites aurantium (as a very abundant
species). Forbes (1848) also dealt with the locality in connec-
tion with Echinosphaerites. Kelly (1860) included the locality
in his Wexford list and, unusually, named some species found
there. Paul (1973) mentioned it, as one of 14 major cystoid
localities, although no details were given. Williams ef al.
(1972:57) noted the place as a shelly locality in the Caradoc
rocks of Wexford and Waterford.

Ballygarvan Bridge, Co. Wexford (1:126720 —
SHEET 23, S792188)

As one of the few fossil localities in south Wexford, Ballyga-
rvan Bridge is often mentioned in connection with Carrigada-
ggan, but similarly until now no systematic re-collection of
the faunas has been attempted since the late 19th century.
M’Coy (1846) listed 9 species; the G.S.I. memoir (Kinahan
1879) has an increased diversity of 11, but with some different
species. The exact locality from which previous collections
were made is in some doubt. Although the description in the
memoir is characteristically imprecise, the G.S.I. 6” field-
sheets have a precisely located asterisk indicating the locality.
However, the geological boundaries adjacent to the bridge
itself are complex and unclear, and since there is no exposure
at the indicated spot the possibility of a cartographer’s
mistake must be kept in mind. The present exposure is very

/BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

10m

WOODEN FENCE 1 ClO $i

ROCK EXPOSURE
CLIFF TOP (ROCK)
VEGETATED SCARP

CONTACT WIT
Oy ERY ING RHYOL

OSED ne
ene

Spe
m

BUSHES & TREES \30/270 = &
Lt a 7 - 63 >
1 ot 7

y¥FROM BASE OF ROCK 7”
1 EXPOSURE IN TUFFS
1 AND SILTSTONES

Pal
YFOSSILS COLLECTED »+++< Ly
3

|Fig.4 Detailed plan of the Kilbride locality.

| limited, but a collection was made from hard grey slates in the
|wooded banks of the river (see Fig. 8), approximately 20m
away along the strike from the supposed locality. The fauna is
sparse, although M’Coy (1846) described most elements of
his list as common. All three collections were probably made
from slightly different horizons in close proximity.

Frankfort and Clogh, Co. Wexford (1:126720 —
SHEET 19, T109550 (Frankfort), T123555
| (Clough))

These are listed by Baily (in Hardman 1887) as one locality.
In fact the 1” sheet 149 and the G.S.I. 6” fieldsheets show
| three separate localities, all of which were investigated in the
|present work. No fossiliferous exposure was found at any
|site, although a few fossils were found in loose blocks at
|Clogh. The localities are the main source of fossils used to
date the Ballymoney Formation of Hiller (1971) and Mitchell
et al. (1972), although Hiller was only able to find wall blocks
of fossiliferous rock, the original G.S.I. localities being
| infilled or overgrown. The localities are shown in Fig. 9.

Raheen, Co. Wexford (1:126720 — SHEET 23,
_$891326)

| The original G.S.I. locality listed by Baily (in Kinahan 1882)
| for 1” Sheet 158 is an old quarry now extremely overgrown,
with almost no exposure. Fortunately, during this study the
local farmer had excavated a new pit roughly along the strike
about 5m away from the old pit, for hardcore. On my
discovery, it was already half filled with domestic refuse and

ie
ea ; N Oo
RUINED COTTAGE ra

(CAT TLESHED) a

ROAD TO TRAMORE ——s

is probably by now completely filled. This temporary expo-
sure allowed collection of a large fauna. Fig. 10 shows the
exact position of Raheen. The locality was not known to
M’Coy (1846) or Kelly (1860), and after collection by the
G.S.I. no attention was paid to it until the 1970s. Shannon
(1979a: 46) recorded a fossil assemblage indicative of a
Caradoc age. Brenchley ef al. (1977) mentioned it in connec-
tion with specimens of Plaesiomys, presumably from existing
collections. It is an important locality, therefore, in that it
provides data along strike from better-known sections at
Enniscorthy, and between there and Carrigadaggan to the
southwest. It is also important in that the newly collected
fauna differs somewhat from previously listed assemblages.

Greenville, Enniscorthy, Co. Wexford (1:126720 —
SHEET 23, S962412)

This locality is the most important of several known from the
environs of Enniscorthy. M’Coy (1846) listed many species
from here, as did the G.S.I. memoir (Kinahan 1882 — locality
5). The locality was reviewed in detail by Brenchley ef al.
(1977), but it was revisited in this study, new material being
collected for the sake of completeness, and with the specific
aim of comparing elements of the brachiopod fauna with
other sampled localities. It was also hoped to collect topo-
typic material of the poorly known agnostid trilobite Trino-
dus agnostiformis M’Coy, the type specimen of which was
redescribed by Whittington (1950: 533). The site location is
shown in Fig. 11. The rock is a very fractured buff coloured
mudstone, occasionally tuffaceous. New material was exca-
vated from shallow depth in the old farmyard, between the

116

Ballyminaun Hill
ae PO

me 0) Kn

Fig. 5 Locality map of the Ballykale locality. Key as in Fig. 2.

lowest doorway of the ruined farmhouse and the new gateway
to the northwest.

Greenville and Moyne Upper Boundary,
Enniscorthy, Co. Wexford (1:126720 - SHEET
23, $967420)

This locality (Fig. 11) should not be confused with Greenville
itself. G.S.I. locality 4 in Kinahan (1882) is Greenville and
Moyne Upper Boundary, where a fauna was re-collected
from many large blocks exposed as a result of very recent
ploughing. These were immediately adjacent to the hedge
bounding the original locality, an old quarry now slurry filled.
No fossiliferous horizon was located within the exposure in
the mainly volcanic quarry.

Courtown localities, Co. Wexford (1:126720 —
SHEET 19, 1187566 (Ballinatray)

Numerous localities in the area of Courtown are listed in an
earlier section, Table 1 (p.110). These were reviewed by
Crimes & Crossley (1968), Brenchley & Treagus (1970) and
Mitchell et al. (1972). Further examination of all these
localities failed, with one exception, to yield anything new or
significant; fossils found were poorly preserved gastropods, a
few external moulds of Glyptorthis and crinoid ossicles from

M. A. PARKES

Gorcy Saw ~

the Courtown Formation. No new material was recovered
from the Ballymoney Formation. The only exception was a
collection made from calcareous slates in the Ballinatray
Formation, about 160 m west of Ballinatray Bridge (Fig. 12).
The present bridge is probably more recent than that named
in the G.S.I. memoir (Hardman 1887), since the Courtown to
Gorey road has been re-aligned since the 6” mapping. There
is considerable confusion about the exact position of localities
in this area, since both the 1” and 6” G.S.I. maps have a

profusion of fossil locality asterisks, not all of which can be |

related to the named localities in the G.S.I. memoir. How-
ever, both from the described position and from the fauna
present it appears that this might be locality 14 of Hardman
(1887), where ‘J/laenus Bowmanni’, ‘Leptaena sericea’ and
‘Orthis calligramma’ were recorded in some abundance.

Slieveroe, Rathdrum, Co. Wicklow (1:126720 —
SHEET 16, 7211890)

This highly fossiliferous locality was known to Griffith’s
collectors (M’Coy 1846) and to the G.S.I. mapping team
(Jukes & Du Noyer 1869). It was re-collected after excava-
tion by Brenchley ef al. (1977) and a mixed brachiopod-
trilobite fauna recorded from there. Although it was
re-collected by me, the combination of very strong deforma-
tion in the fragile slates and shortage of time meant that no

|
|

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

lo fe.
‘S
A

| WZ, HF fossil lgcali

CLOLOGE UPPER

|
Fig. 6 Locality map of the Clologe Upper locality. Key as in Fig. 2.
|

further work was done on the material. The previous faunal
lists are used in discussion of correlations.

Other localities

In the course of tracing the localities from the memoirs,
several other records of fossils were noted. On Sheet 158
(Kinahan 1882: 26) a locality called Kellystown Bridge was
described a little south of Raheen, but no details were given
in the palaeontological notes and it was not traced on the
G.S.I. 6" fieldsheets nor in reconnaissance fieldwork. Kelly
(1860) gave details of various localities which were not all
included on the G.S.I. 6” sheets or in the memoirs, including
‘Ballyminaun Hill and Carriganeagh, both south of Gorey.
‘These do not now have any fossiliferous exposure. Fossils
were recorded by Shannon (1979a) from Raheen and Bally-
brennan, both G.S.I. localities, and also from Wilton Castle
southwest of Enniscorthy, but the area is now heavily for-
ested and no fossils were found in the restricted exposure
available. All other sites examined as a result of personal
ommunications, or unnamed sites indicated by asterisks on
1” and 6" geological maps, proved to be either unfossiliferous
: not now exposed.

117

A \W oS
O
ALS PY

METHODOLOGY AND TECHNIQUES

Sampling methods

In all sampled localities, an initial collection was made by
identification of the fossiliferous lithology and on-site collec-
tion of specimens. Subsequently, the main collection of
specimens was achieved by removing large volumes (between
10 kg and 80 kg, occasionally more) of the fossiliferous
lithology, to be broken up and examined in a laboratory. The
validity of this method was verified by the fact that successive
seasons’ collections served to increase the numbers of speci-
mens, but not the diversity of the fauna. The one exception to
this was the Kilbride locality near Tramore, Co. Waterford
(p.112). This debris flow appears to have ‘sampled’ various
benthic associations in its downslope movement, and subse-
quent collections made here increased the diversity of the
fauna significantly from the initial sampling.

118

fossil

locality

Fig. 7 Locality map of the Carrigadaggan locality. Key as in Fig. 2.

Preparation techniques

Standard preparation methods were used. The measurements
of all material (in millimetres) were made using vernier scale
calipers or a micrometric graduated microscope eyepiece,
both accurate to 0-1 mm. The combination of camera, lenses
and extension rings (Table 5) gave a range of magnifications
of up to X5. The specimens were whitened with ammonium
chloride sublimate before being photographed.

Taxonomy and statistical analysis

In this study both multivariate and bivariate analyses have
been utilized where the measurement data were adequate.

Table 5 Magnifications obtained by different lens and extension
ring combinations.

AF Micro-Nikkor 55 mm f/28 x1
AF Micro-Nikkor 55 mm f/2-8, PK11A, PK12, PK13 <<?
24 mm lens reversed + PK11A x3
24 mm lens reversed + PK13 x4
24 mm lens reversed + PK11A + PK12 + PK13 x5

NOTE: All other combinations produce non-integer magnifications.
Magnification less than X1 necessitated the use of the AF Micro
Nikkor 55 mm, but not fully extended. The magnification was
calculated by comparison of the negative with the measured
specimen size.

M. A. PARKES

arrickbyrne

Hill

For detailed discussion of statistical methodology in brachio-
pod systematic work, reference can be made to Williams
(1962), Harper (1984) and Temple (1987). In many older
palaeontological references the use of statistics to define
species is minimal or non-existent. For example, the funda-
mental work of M’Coy (1846) quotes the length of most
species described as a single value (in inches and lines, where
a line = 1/12'" of an inch). No indication is given as to
whether this is the length of a figured specimen or a subjec-
tive assessment of the mean length of a sample. Here a
pragmatic approach was taken, and measurements were
made on any specimens worth measuring. Some measure-
ments taken on slightly deformed material are included in
descriptions in the text as a guide to proportions, but where
this occurs it is pointed out. Table 6 is a set of defined variates
as used by Harper (1984) and adhered to in the present work.
Although in some cases the measurements taken did not
provide sufficient reason to classify a sample as belonging to a
particular species, the original data for these and all mea-
sured specimens is lodged in the General Library Biological
Data Collection at The Natural History Museum, South
Kensington, London. The figured specimens themselves are
deposited in the Palaeontology Department collections there,
register number prefix BC. Another set of the data is lodged
in the James Mitchell Museum, University College, Galway,
where all non-figured brachiopods and other material are
deposited. Analysis was done using the ‘Palstat’ package on
BBC microcomputers (Harper & Ryan 1987).

| BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

STRATIGRAPHICAL CORRELATION
|

The stratigraphy of SE Ireland, as presently understood, is
the result of many years work by innumerable individuals.
The present study is concerned only with one part of the
“succession — the Duncannon Group, which is a mid-Upper
Ordovician (mainly Caradoc) volcanic-dominated sequence,
with clastic sedimentary intercalations. It is generally uncon-
formable on the Lower Ordovician (Arenig — Llanvirn/
Llandeilo?) Ribband Group.

In certain better-exposed areas, notably Courtown
(Brenchley & Treagus 1970, Mitchell et al. 1972) and
_ Tramore (Stillman 1976, Carlisle 1979), more detailed strati-

graphies have been erected with several local formations. The
wo areas of Courtown and Tramore were correlated on the
asis of faunas and comparable stratigraphies by Mitchell et
al. (1972), and the age subsequently revised by Carlisle

1979) through better knowledge of the extent of the gracilis

ne. Thus both the lowest calcareous horizons, the Tramore
Limestone Formation and the Courtown Formation, are
-onsidered as late Llandeilo in age, or even older (Llanvirn-
iim ine based on conodonts (Bergstr6m 1971).

The thick development of volcanics west of Tramore and
aterford is perhaps the most complex area of the Duncan-
ion Group and there is much debate as to the true sequence

Fig. 8 Locality map of the Ballygarvan Bridge locality. Key as in Fig. 2.

119

there. Tietzsch—-Tyler (1989) reviews the various arguments
and draws his own interpretations for the G.S.I. compilation
maps used in Parkes (1990). Stillman (1976) has noted that
the succession is subdivided on lithostratigraphical grounds
and should be regarded as comprising ‘volcanogenic units’,
essentially derived from a volcanic centre situated about 4 km
northwest of Tramore, and another farther west. The impor-
tance of fossil assemblages to date horizons within such
sequences is thus clear, to confirm and supplement the
petrological and field relationships of the units. The Tramore
Limestone Formation and some gracilis-bearing shales were
the only well-dated horizons until Carlisle (1979) recorded a
shelly assemblage from Kilbride, which is described herein
(Table 11, p.129). The assemblage, near the top of the
succession, provides some constraint on the date of cessation
of volcanism in Waterford. Consequently, although the age
of the Kilbride fauna is discussed below, the remainder of the
sequence and its faunas underlying the Kilbride assemblage is
outside the scope of this review, as previously noted.
Re-examination of the Courtown localities collected by
Crimes & Crossley (1968), Hiller (1971), Brenchley & Trea-
gus (1970) and Mitchell et al. (1972) yielded no significant
new information and the formations erected by these authors
and the existing age constraints described from the fossil
assemblages are thus accepted here. The development of

ime

NF
ies 3

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

Fig. 10 Locality map of the Raheen locality. Key as in Fig. 2.
| Table6 Definition of variates measured on the fossil brachiopods
in this paper.

sagittal length
X2 maximum width
X3 so position of maximum width measured from posterior margin
maximum depth measured perpendicular to sagittal length
hinge width
maximum length of interarea
_ X7 origin of fold or sulcus measured from posterior sagittal
margin
X8 maximum width of rim or limbus
X9 maximum length of muscle scar measured from posterior
sagittal margin
X10 maximum width of muscle scar
|X11_ wavelength of median rib at 5 mm growth stage
position of apex measured anteriorly from posterior margin
X13 length of base of spondylium
X14 maximum width of spondylium
length of base of cruralium
X16 = maximum width of cruralium
|X17 anterior extension of brachiophore bases measured from
umbo in plane of sagittal length
transverse separation of ends in brachiophore bases
X19 maximum depth of sulcus
maximum width of sulcus
X21 length of cardinal process base
X22 sagittal length of submedian septa
X23 __ transverse separation of anterior ends of submedian septa
X24 sagittal length of median septum

calcareous clastics followed by black graptolitic shales within
gracilis times appears to have been restricted to the areas
mentioned above, since no comparable sequence is recog-
|

121

RAHEEN

fossil locality

nized elsewhere in the Duncannon Group, and has not been
found in boreholes.

In between these two areas, the main belt of Duncannon
Group rocks is poorly exposed, often as resistant hills of
rhyolitic intrusions or thicker volcanic units which were
probably original volcanic centres. The succession is poorly
differentiated, despite several intensive studies. The Duncan-
non Group was defined by Gardiner (1974) from the Duncan-
non district of southwest Wexford and divided into four
formations; Duncannon Volcanic Formation, Arthurstown
Formation, Ballyhack Formation and Campile Volcanic For-
mation, all of Caradoc age based on graptolite evidence, in
the regional Campile Syncline (Gardiner 1970). However,
Shannon (1978, 1979a), in mapping the area south of Ennis-
corthy, but not continuous with the area of Gardiner (1967),
differentiated the succession into only two Formations: the
Doonooney Formation (with the Ballybrennan Volcanic
Member) and the younger Raheenahoon Volcanic Forma-
tion, which encompasses the main rhyolitic volcanism. Both
Raheen and Carrigadaggan are within the latter formation in
the area mapped by Shannon. The remapping of mid Co.
Wexford (Geraghty 1989) north of Shannon’s area has con-
firmed the threefold division into Cahore, Ribband and
Duncannon Groups.

Despite the poor exposure and limited extent of the inlier
at Kildare, the sedimentary succession is more varied and the
palaeontological constraints on the age of different horizons
is better understood, although there are many problems
remaining. The Duncannon Group equivalents probably lie
unconformably upon the older rocks below them. A Soud-
leyan fauna and a Longvillian fauna constrain the age of
volcanism here, being found above and below the lavas of
Grange Hill. There are Ashgill faunas in the Kildare Lime-

|
_ BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

| Fig. 12 Locality map of the Ballinatray locality. Key as in Fig. 2.

| stone Formation (Rawtheyan) and the well known Hirnantia
fauna in the topmost mudstones. A review of this inlier is
given by Parkes & Palmer (1994). The stratigraphy of the
| eastern Ireland successions in the Iapetus suture zone has
been investigated by Murphy (1987), who identified two
' separate terranes, the Grangegeeth and Bellewstown ter-
ranes, between the Balbriggan-Herbertstown sector of the
Leinster terrane and the Central terrane (equivalent of the
| Southern Uplands of Scotland). These terranes are identified
on the basis of contrasting volcanic characteristics and strati-
graphical sequences as well as faunal differences. Murphy
(1987) also correlated the sequences of eastern Ireland with
those of the Leinster terrane and the Lake District of
England.
Within the Leinster terrane, recent detailed reviews of the
| complete stratigraphical successions have been by Williams et
val. (1972), Briick et al. (1978, 1979), and Holland (1981).
More recently Harper & Parkes (1989) summarized the
palaeontological constraintson the development of Irish Cale-
donide terranes, a subject expanded upon in Murphy et al.
(1991). Parkes & Vaughan (1992) and Owen et al. (1992)
have dealt specifically with the Grangegeeth terrane.
Tietzsch-Tyler (1989) has completed a re-evaluation of the
Lower Palaeozoic geology, as well as the preparation of
G.S.I. compilation maps, for south-east Ireland. This latter
work involves numerous modifications which are outside the
Scope of the present work, but several redefined formations
are used in the Duncannon Group, as explained in the key to
those maps. Principal of these is the Annestown Formation,
of essentially rhyolitic composition with andesite, slate and
limestone members. This represents undifferentiated middle
Duncannon Group sediments and volcanics on the map, and
is the equivalent of the Doonooney and Raheenahoon Volca-

: ~
atray Bridge
> ee

123

>
|
>
|
>
yd
ey

nic Formations of Shannon (1978). It is also equivalent to the
Lower Tramore Volcanic Formation and all formations
above (except the intrusive rhyolites in the Upper Tramore
Volcanic Formation of Carlisle (1979)), the Duncannon,
Arthurstown, Ballyhack and Campile formations of Gardiner
(1974), and the Ballymoney and Gorey Rhyolite formations
of Brenchley & Treagus (1970), as well as the Upper Caradoc
Raheen Formation of Owen et al. (1986). Tietzsch-Tyler’s
Carrighalia Formation (equivalent to the Ross and Loftusa-
cre Formations) with its Tramore Limestone Member is also
the equivalent of the Courtown and Ballinatray formations of
Brenchley & Treagus (1970).

AGE OF THE ASSEMBLAGES

Kildare, Grange Hill, Horizons 1 and 2

Wright (1970) claimed that these were Longvillian in age, on
the grounds that their composition is similar to assemblages
from the Gelli-grin Formation of the Bala district of Wales,
and in particular in relation to the presence of the calcareous
inarticulate Orthisocrania divaricata, whose occurrence could
only be substantiated in Longvillian rocks in Britain.

Whilst there are some differences noted (Tables 8, 9,
p.128) between the assemblage described herein and that
listed by Wright (1970), in essence the fauna still shows a
strong affinity with the Welsh Bala faunas described by
Williams (1963). Many of the genera are long-ranging in the
Caradoc, or even the Ordovician, and where samples are
inadequate to determine the species, then the overall generic
composition has been used. However, at Grange Hill both

124

Orthisocrania divaricata and Cremnorthis parva occur, and
these are at present only known from Longvillian rocks in
Britain. Leptestiina oepiki is also recorded first in Longvillian
Welsh localities (Williams 1963). Although Cremnorthis
(Hints 1968, Llandeilo) and Orthisocrania (see p.136) are
known from older Baltic successions and may have migrated
to Leinster at a time different from their Welsh Basin arrival,
on balance, with the total assemblage affinities with the
Gelli-grin Formation, a Longvillian age is most likely for this
assemblage.

Given the close relationship of Grange Hill Horizon 2 to
Horizon 1 with a substantially similar but reduced diversity
assemblage, a similar Longvillian age is suggested for this
horizon also. The position of the various localities at Kildare
in the stratigraphical succession of the inlier is schematically
illustrated in Fig. 13.

Kildare, Grange Hill House Cottage

The only modern reference to the faunal assemblage from
this locality is Williams ef al. (1972). A Soudleyan age is
claimed for the fauna, presumably on the basis of compari-
sons with the faunal list published by Reynolds & Gardiner
(1896). The recollection of the fauna, as described herein,
confirms that suggested age. The palaeoecology of this
assemblage differs substantially from other Duncannon
Group faunas described here (Table 10, p.129), but the
brachiopod species show a strong similarity with a fauna from
Herbertstown, in the northernmost part of the Leinster

green
sandstones

SILURIAN

| _red mudst. |
black shales
HIRN. mudstones

Kildare
Limestone Fm.

GRANGE
HILL
HOUSE
mudstones COTTAGE

sandstones

eo)
=
Lu
a)
z
<
-!
=)
z
oc
>
Zz
<
a
_

ARENIG

Fig. 13 Schematic section of the succession in the Kildare inlier.

M. A. PARKES

terrane, described by Harper et al. (1985). The combination
of Plaesiomys multiplicata and Oanduporella cf. reticulata is
unknown elsewhere in the Leinster terrane (except possibly
at Clologe Upper). Indeed, the enteletoidean Oanduporella
was unknown outside the Baltic before the Herbertstown
record. The assemblage at Herbertstown was noted as being
unlike any other fauna recorded in eastern Ireland (Harper et
al. 1985: 289), but the new Kildare sample shows strong
resemblances. On balance, the Herbertstown fauna was
restricted to the Caradoc, with a most probable Soudleyan
age. Similarly at Kildare, the fauna cannot be younger than
Longvillian since it lies below the andesites of Grange Hill,
which themselves are below a Longvillian assemblage. On the
whole a Soudleyan age is most likely since the common
brachiopod Plaesiomys multiplicata is found in the Soudleyan
of Glyn Ceiriog of North Wales (Bancroft 1945) in associa-
tion with Rafinesquina, which is present at Kildare too. The
relationship of this horizon to others at Kildare is shown in
leg, I}.

Kilbride

At this locality Carlisle (1979) listed a fauna with some forms
congeneric with those in the fauna described here (Table 11,
p.129). There are several differences, but the principal gen-
era are all indicative of a Caradoc age. Carlisle (1979: 552)
recorded Decordinaspis, a trinucleid trilobite previously only
found in the Harnagian/Soudleyan of Grangegeeth (Harper
& Romano 1967). Thus she suggested a Soudleyan age for the
base of the Upper Tramore Volcanic Formation, in which
Kilbride is located, with an unknown upper limit but possibly
extending into the Upper Caradoc. This author did not
identify Decordinaspis here, but, based on the co-occurrence
of Cremnorthis parva, Leptestiina oepiki and Sowerbyella
sericea, a Longvillian age is postulated for the assemblage.
This accords well with Carlisle’s interpretation. If the Orthis-
ocrania she recorded proves to be O. divaricata, this would
further enhance the reliability of the age assigned, since the
locality was said (Carlisle 1979: 551) to be 200 m above the
base of the formation.

Ballykale

This assemblage is relatively low in both specimen numbers
and diversity, being dominated by Bimuria cf. dyfiensis
(81%, see Table 12, p.130). This species has only been
described previously from the Gelli-grin Formation of north
Wales (Lockley 1980:215), which is of Longvillian age. In the
other localities dealt with here, Bimuria sp. definitely occurs
only at Kilbride, in a form probably conspecific or at least
very close to B. cf. dyfiensis. This locality too is of probable
Longvillian age and this seems the most reasonable estimate
for the age of the Ballykale assemblage also. The few other
brachiopods are all congeneric with Kilbride forms, notably
the rare Ptychoglyptus.

Clologe Upper

Since the poor preservation of fossils from here permitted
only a generic level identification at best, the precision of
correlation can only be crude. Overall, the generic composi-
tion indicates a Caradoc age and some strong similarities with
the other assemblages described herein. However, the pos-
sible occurrence of both Oanduporella and Plaesiomys, both

== ag — axe .

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

reminiscent of the Soudleyan fauna at Grange Hill House
Cottage, Kildare and the Soudleyan Herbertstown assem-
blage (Harper et al. 1985), together with the apparent
| absence of Cremnorthis and Orthisocrania, suggest that the
| assemblage is older than Longvillian. Therefore, a probable
Soudleyan, or perhaps Harnagian, age is postulated.

Carrigadaggan

| The brachiopod fauna contains numerous elements which
_ allow confident assignment of the assemblage to the Longvil-
lian, and enhance the strong inter-locality correlation. Lep-
testiina oepiki, Cremnorthis parva, Orthisocrania divaricata,
| Sowerbyella sericea, Skenidioides costatus and Kullervo aff.
hibernica are all found in the Longvillian of Grange Hill,
Kildare, as are many of the genera such as Nicolella,
Platystrophia and ‘Orthambonites’. The species listed above,
except Orthisocrania and Kullervo, are here recorded at
Kilbride also (Table 14, p.131); Carlisle (1979) also listed
_ Orthisocrania from there. The presence of Saukrodictya sp. is
noted here, which although rare, also occurs at Kilbride,
| Ballykale and Greenville-Moyne, all of which are probably of
Longvillian age. The cystoid Echinosphaerites ct. granulatus
is most common here, but was also found at Kilbride and
| Ballygarvan Bridge. Although not stratigraphically useful, its
occurrence in these three geographically close localities is
| supportive of the brachiopod correlation.

_Ballygarvan Bridge

The sparse brachiopods from this locality allow little certainty
_about correlation of the assemblage, neither does the
restricted trilobite material. The generic composition and
abundance of gastropods (Table 15, p.131) is notably similar
to that of Grange Hill (Table 8, p.128) and Grange Hill
House Cottage (Table 9, p.128) at Kildare. This may be
| merely ecological control. However, as the large inarticulate
Lingulella ovata occurs elsewhere in Leinster only in the
| Longvillian of Grange Hill, Kildare, a possible Longvillian
age is suggested. In spite of the low numbers of specimens,
the congeneric forms enhance the consistency of inter-locality
correlation in the Duncannon Group.

‘F rankfort and Clogh

The examination of these localities was unproductive for new
_assemblages. However, the collections made by N. Hiller (for
‘a B.Sc. thesis, 1971) and subsequent publication (Mitchell er
jal. 1972) correlating the Courtown area with the Tramore
region, and now housed in the Ulster Museum in Belfast,
| were examined for comparison with other collections made in
the present study. The list of genera published by Mitchell et
al. (1972) is quite long but does not reflect the proportions
within the assemblage. The strong deformation and poor
preservation in the tuffaceous rocks, collected from walls in
\the area of the G.S.I. locality, made unequivocal identifica-
|tions difficult. Many genera are represented by one question-
able specimen only. The abundant genera are Cremnorthis,
‘Orthambonites’, Leptestiina and dalmanelloids. Less com-
‘mon but positively present are Kullervo, Platystrophia,
Anisopleurella, Skenidioides, Nicolella and Orthisocrania.
The apparent absence of some of the other listed genera
could be due to incomplete donation of the collection, rather
than absence from the assemblage, since not all specimens

125

were clearly labelled. The particular elements not seen in the
material at the Ulster Museum were Diambonia, Ptychoglyp-
tus, Eoplectodonta, Pseudolingula and Actinomena? The
Christiania recorded is a somewhat equivocal single specimen
which is highly strained.

Thus, if the definitely present and abundant elements alone
are considered, the assemblage clearly takes on a strong
resemblance to the described assemblages from Grange Hill,
Carrigadaggan, Greenville-Moyne and other localities in
Wexford. The re-examination of this assemblage, in conjunc-
tion with a revision of the Duncannon Group localities
throughout Wexford and elsewhere in the Leinster terrane,
reinforces the Upper Soudleyan — Longvillian age assigned to
the Ballymoney Formation by Carlisle (1979: 552; fig. 3). On
the basis of the existing brachiopod collections this will not be
refined further, but new material or the current examination
of the trilobite collections in the G.S.I. may yield better
information.

Raheen

The assemblage described here (Table 16, p.131) is clearly
inadequate for a confident correlation and is probably facies
controlled, rather than of different age from other localities
in the Duncannon Group. When the trilobite identification is
complete a better constraint on the age might be made, but in
the absence of better material the G.S.I. lists (Kinahan 1882)
give a good indication of the age of adjacent beds in the
succession. There are no species known to be restricted
temporally, but typical Longvillian genera are absent. Some
elements are reminiscent both of Greenville and the Brick-
works Quarry Shale Formation of Grangegeeth (Romano
1980a), which are believed to be Harnagian or Soudleyan.
Therefore, an early Caradoc age, possibly Harnagian or
Soudleyan, is thought to be the best estimate.

Greenville

The collections made here (Table 17, p.132) are somewhat
reduced by comparison with those of Brenchley ef al. (1977).
These authors suggested a Harnagian age, and no significant
evidence has been found to dispute that in the assemblage
described here. Only the probable occurrence of Oandupor-
ella, present in the Soudleyan of Herbertstown (Harper et al.
1985) and in the Soudleyan assemblage at Grange Hill House
Cottage, Kildare, suggests a possible Soudleyan age. Brench-
ley et al. (1977) attached some significance to the presence of
‘an early species of Sericoidea together with Anisopleurella
aff. multiseptata Williams’. As discussed in the systematic
description of Chonetoidea (the senior synonym of Seri-
coidea), the variation amongst the population is considerable
and assignment to C. aff. abdita Williams would now only
suggest a Lower Caradoc age. Anisopleurella multiseptata,
described from the Costonian Derfel Limestone Formation
(Whittington & Williams 1955), is known also from the
Longvillian Gelli-grin Formation of Bala, North Wales
(Lockley 1980).

Greenville-Moyne

Several elements, such as Cremnorthis parva, Leptestiina
oepiki and Skenidioides costatus, are conspecific with samples
from Longvillian assemblages at Kilbride, Carrigadaggan and
Grange Hill Horizon 1 at Kildare. Other forms only identifi-

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dnoiy
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snjnosnjese}

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ueIyOOsqusueyy
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ANOZ 3LNOLdVYS

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

' able generically or questionably assigned, e.g. Nicolella sp.,
_‘Orthambonites’ sp., Orthisocrania?, Rhactorthis? and Sower-
| byella?, are all reminiscent of those assemblages, as is the
' rare occurrence of Saukrodictya sp. Kullervo aff. hibernica is
‘also present at Greenville-Moyne, Carrigadaggan and
Grange Hill Horizon 1, all localities thought to be Longvillian
in age, but it also occurs in the Knockerk House Sandstone
Member (Romano 1980a) of the Grangegeeth Group
(equivalent to the Upper Tuffs and Shales of Harper, 1952),
_which is believed to be Costonian in age (Brenchley et al.
1977). However, the consistency of the inter-locality correla-
| tions suggest that a Longvillian age is most probable for the
_Greenville-Moyne assemblage.

Ballinatray

Although the assemblage from here is too deformed and
broken to identify the brachiopods precisely, the age of the
horizon is constrained by previous work. The age of the older
Courtown Limestone Formation is believed to be mainly
Llandeilo ranging up to the Costonian Stage of the Caradoc
on the correlation with the Tramore Limestone Formation
(Carlisle 1979). The succeeding Ballymoney Formation is
believed to be of probable Longvillian age, and in any case
gracilis Biozone graptolites have been recovered from the
Ballinatray Formation (Mitchell et a/. 1972). An early Cara-
‘doc, at least partly Costonian age is suggested for the
\formation.

In the light of the age constraints clarified by a revision of
these Duncannon Group faunas a revised correlation chart
has been produced (Fig. 14), with sections additional to
previously published correlations e.g. Williams er al. (1972)
and Briick et al. (1979).

PALAEOECOLOGY AND BIOGEOGRAPHY

|
|
|
|
|
|

In some of the localities very little ecological data can be
derived because the exposures were small, resricted and
isolated from a clearly defined succession, or assemblages
were not from in situ rocks. Equally, small assemblages like
those from Ballygarvan Bridge (Table 15) are inadequate for
detailed interpretation. Even with large assemblages such as
Grange Hill Horizon 1 (N=646) some reservations are held
about interpreting too much from them.

| One evident point from this work is that assemblages often
differ from previously recorded collections, especially those
of M’Coy (1846) and the G.S.I. Memoirs. Many differences
are simply taxonomic artefacts reflecting better systematic
differentiation of this important phylum. Where the outstand-
ing differences are not addressed directly below, the best
explanation is that in the volcano-sedimentary setting of these
ocalities the character of faunas changed as a result of tuff
falls etc. and slightly different horizons have been sampled.
This would apply to Ballygarvan Bridge, Raheen, Greenville
and possibly Greenville-Moyne for example.

For each locality a table of the total collected fauna is
resented. Table 7, however, is a summary chart of presence/
absence data for each locality, highlighting the inter-locality
affinities. The tables (8-19) include total counts of the
brachiopod valves and various animals for the comparison of
assemblages with recognised palaeocommunities discussed
pelow. Following Lockley (1980: 171) this number has been

127

Table 7 Summary chart of fauna collected at each locality.
Localties: A — Kildare, Grange Hill Horizon 1. B — Kildare,
Grange Hill Horizon 2. C — Kildare, Grange Hill House Cottage.
D — Kilbride. E — Ballykale. F— Clologe Upper. G -
Carrigadaggan. H — Ballygarvan Bridge. I— Raheen. J -
Greenville. K — Greenville-Moyne. L — Ballinatray.

MJ} CJD I FOG isi tl Y IK Ie

BRACHIOPODS
Acanthocrania? OS tS 70] 5S 6 ee
Anisopleurella cf. multi-

septata = Cy e e -
Bicuspina?
Bimuria cf. dyfiensis ~ e =
Bimuria sp.
Chonetoidea abdita
Cremnorthis parva
dalmanellids indet.
Dolerothis? - -
Glyptorthis a
Hedstroemina sp. Ce SE nS ee
Hesperorthis sp. ere es lee eee Oe sy
Hibernodonta? Sh Oe i te ee fae —
Kjaerina? e =
Kjerulfina? e -
Kullervo aff. hibernica == = = Ser = Se =
Leptaena sp. e

Leptellina cf. llandeiloensis —
Leptestiina oepiki e--eee
Leptestiina oepikiampla —
Lingulella ovata ee -
Nicolella cf. actoniae e
Oanduporella cf. reticulata —
‘Orthambonites’ sp. e
Orthisocrania divaricata @
Oxoplecia?
Petrocrania harperi
Philhedra sp.
Plaesiomys multiplicata —- -— @ — —
Platystrophia sp. O46 =. @ Te = geneh= = ee
Porambonites sp. - - e

Ptychoglyptus sp. =—— -—_— 10) (0) =) Se
Rafinesquina sp. - - @®@------ -
Rhactorthis sp. e- -
Salopia sp. - @ -
Saukrodictya cf. sp. A -- -
Schizotreta sp. -
Skenidioides costatus e
Sowerbyella sericea e
Strophomena? e
TRILOBITES
Ampyx austinii - - @®----- @®e- -
Arthrorhachis e
Atractopyge =SSseae es OS sas =
Autoloxolichas e - -
Calyptaulax e----- @---- =
Deacybele - e
Encrinuroides - = - = = = = = =
Flexicalymene e e

Homalopteon sp. - =
Remopleurides -------- -
Tretaspis ----- @-- @®- - -
‘Trinodus agnostiformis’ — — — — e
Xylabion e - - - - =
OTHER ELEMENTS
Echinosphaerites
granulatus
cheirocrinid cystoid
crinoid ossicles
gastropods

ramose bryozoans
prasoporid bryozoans
tentaculitids
orthocones

bivalves

conulariids

ostracods

eee!
il
|
leoe0ee
| e |
eee!!!
eeoec0c!|
e
e
eeo0e
eee!
ee!

ee!
e |
|
e |

eeeeeoeeee i!
e |
|

eee!

ee!

|
|

128
calculated using the formula:
N=A + 0-51 + P (if P>B), or N = A + 0-SI + B (if B>P)

where: A = no. of articulated valves, B = no. of brachial
valves, P = no. of pedicle valves, I = no. of indeterminate
valves.

The problems of counting different groups of animals was
discussed by Lockley (1980: 171-2) for Welsh Caradoc faunal
associations and by Jaannusson (1984) for Baltoscandian
Ordovician sequences. In the case of groups other than
brachiopods absolute numbers are given. However, for some
organisms such as fragmentary bryozoans, cystoid plates and
crinoid ossicles, absolute numbers are given in brackets and
an equivalent number of individuals given, generally one. An
arbitrary correction factor is applied to bivalves for Kildare
Grange Hill House Cottage, by halving the total, on the
assumption that the valves are simply disarticulated in equal
proportions. This is also applied to ostracods. For trilobites
the largest value of pygidia or cephala is arbitrarily taken as

Table 8 Total assemblage collected from Kildare, Grange Hill
Horizon 1.

Internal External Total
Pw Bw © i PY C i wo, YW

BRACHIOPODS
Acanthocrania? | tee = 1 Sa It @eils)
Chon. toidea sp. 1 <= 18=9.]— le si 2035
Cremnorthis parva St WH 4 = 1OAl SF = 5 37
dalmanellids, indet. 23) = 3S 8) 2 hy ANP)
Hedstroemina sp. SSS eS 3 = 3 OS
Kjerulfina? PSS Sa = = = i Pil
Kullervo aff. hibernica 28-<=8=8 1 = =PS9 2203
Leptaena sp. Goi oa 35 =] 6 i tks
Leptestiina oepiki 7 === = = isa elO
Lingulella ovata - -=-- =-=-- 3 3 055
Nicolella cf. actoniae = 1 = SST = S05
‘Orthambonites’ sp. 256 =8—) 20 = = = aorn0:9
Orthisocrania divaricata 38 23 —- 68 -— - - 108 92 14-2
Petrocrania harperi = 89 = ==—30 = = 89) 13:8
Philhedra sp. =8§=8—8=) — Ieee eles
Platystrophia sp. Sl EG = We iyi 1 Sy Ss
Rhactorthis sp. 1 <3 ty 1 27 SSO: 8
Skenidioides costatus Wo 32 i = Wilh 3 = Ils ils
Sowerbyella sericea A Wy tl = IPI it @ 38 Sell
Strophomena? 28=-s = 1S = = 2803
TRILOBITES
ANH MUOTOUWGITS CHONG soccoscaenncadqnnnsne: oonnandABSnDS0on0006 3) eS)
Cally ares? OSGI cooecoasansaqncotnoocosbon0onDooang0500090000 iL sis)
Hlexicaly mene cephalonwerrce--eeerer eee eeeeeeee errr crete ee raee i pil
OTHER ELEMENTS
ClatsXorralvoxsyq io tox (7) gaepaeatoondpabEdnacaocotsdagedeenoswapedoqcebe iL (Opil5)
PASOPOUS ee coer mrceckiecses. dee iaeeeemeeasccoeesaciewecsineet S30) Fed
TAMOSE DLV OZOANS meme eree eeeece eee eee eee eeeeeac eer ere 4 0-6
prasoponidibiyOZOanSmen settee eee eee eee ee eee eee eee eee ile
tentaculitids: .eecmaceesmesserecac een chisa tclarcosecenelekee sites 29 =4-5
OnthOCONES resesice chee socGeedeereh se eeeee alee meee ener 4 0-6
BIVAlVES: Aiciccsecgemetee de egetcinevseueseneerntecsee soak seeliemtaeee act 2:3)
COND ALIGS’. «ca. estan gee neh acteesigs soeeme eee aeeepecseesseceaeses i pil
SPONGESPICUIE He ctewesceee am atradeossemeresee ts -eecles ceeds. il. OpitS)
Total 646 99-85

PV = pedicle valves, BV = brachial valves, C = conjoined valves, I =
indeterminate valves, NO. = equivalent number of total animals. (N.B.
Numbers in brackets are total fragments of colonial or fragmented individuals
— see discussion). See discussion on p.127.

M. A. PARKES

the commonest animals, errors involved in counting other —
animals are considered as negligible.

The diverse assemblage from Kildare Grange Hill Horizon
1 is listed in Table 8. All the material consists of moulds from |
decalcified mudstone/siltstone. The ratio of ventral to dorsal |
valves is generally equal. Differences may be accounted for —
by the problems of identifying fragments produced by the |
sampling method. There is a large range of size variation |
between and within species, and there is no suggestion of
current winnowing. The shells are disseminated through the
sediment, not collected together as in a coquina. For these
reasons the assemblage is treated as a palaeocommunity.
Grange Hill Horizon 2 (Table 9) is very similar but has
reduced frequency of fossils and the addition of common —
Lingulella ovata. The difference is partly because the rock is
not decalcified. This biased the identifications in favour of
larger distinctive species such as L. ovata, O. divaricata and
Platystrophia. Another factor is that at least one thin tuff
horizon is found between the two horizons, which may have
subtly changed the assemblage.

Some differences between Table 8 and the faunal list of |
Wright (1970) should be noted. Cyclospira and Bicuspina
were not collected by me, whilst taxa not recorded by Wright |
but collected by me include Acanthocrania?, Chonetoidea,
Hedstroemina, Kjerulfina, Kullervo aff. hibernica, Leptestiina |
oepiki, Lingulella ovata, Nicolella ct. actoniae, Petrocrania |

}
|

q
|
|
|
the number of individuals. Since brachiopods are generally |
|

harperi and Philhedra. All these were present as 1% or less of
the assemblage, except for Petrocrania which alone consti-
tuted 13-8% of the assemblage. Wright (1970) also recorded
Eoplectodonta, but this genus is essentially indistinguishable |
from Sowerbyella except for the presence of denticles along |
the hinge line. No specimens in this study showed evidence of |
such denticles.

The older fauna from Grange Hill House Cottage (Table
10) is somewhat anomalous in comparison with the other

Table 9 Total assemblage collected from Kildare, Grange Hill
Horizon 2.

Internal External Total
VIB (CU WAY EAC It no. %
BRACHIOPODS
dalmanellids, indet. 2n3x=)= os thin meee Sail
Hedstroemina sp. 7 --- Li==.5, 1 11-9
Leptaena? Sa Le 3-3
Lingulella ovata - -1- ---16 9 153
Orthisocrania divaricata erage ey a) 8:5
Petrocrania harperi 2 4 - 4 6-8
Platystrophia sp. S)ieov=y Sh2e= silgas B)|
Salopia? Spliey a—) 1 17
Skenidioides costatus 3 --- --- - 3 Sal
Sowerbyella sericea 2 - - 2 333
OTHER ELEMENTS
@) EU NVOYHO) 1S Gagan goadcnan ance sodecadoogase sueagaRaDoIadacccooGaaZb0ue 3 5-1
[DHAO) KOVAC! lOVAYOVAOHIN scceosndancsoepsoobe ssbocodenapsecopeesconan 3) Sal
DivalVe: sce dioscenee cht. teats seat ea- te netbe Steels ances epee CER 1 1-7
(ACHR) SXOTEES, densesuedassondsne sasaccbrcsoee cose dgubbonecBosoncSsBCHsc5 13 22:0
Total 59 100-0

SEE

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

Table 10 Total assemblage collected from Kildare, Grange Hill
House Cottage.

Internal External Total
PVBVC I PVBVC I no. %

~BRACHIOPODS
dalmanellids, indet. = ate SW IS 2 0-4
Dolerorthis? SS SS 1 0-2
_ Hibernodonta? = 3 == 2 = 3 4 0-8
_ Leangella? 1 = 1 0-2
Leptaena sp. fe= = Sse s 1 0-2
lingulid, indet. ==S=s5 =+=+-+4 4 0-8
Oanduporella cf. reticulata 13 10 — 9 6-- 13 2-5
| Petrocrania? el 2 0-4
| Plaesiomys multiplicata 2613 —- — 12 20 — - 26 5-0
Rafinesquina sp. § V= 2 10 Ss = 4 12 2-3
Skenidioides sp. 3. = 5) Sages 3 0-6

TRILOBITES
_ Flexicalymene QDs [OWGRGTIAY Gobasedesaneseconseubeannce neepeas000 2 0-4
| Flexicalymene sp. cephala .............cssserceceesenseneeees 4 0-8
PEO COUGHOSIGEpNAlal .2...o.. sacle sae sire sscijeoane se series i 3 0-6
EIPIGSP: PYSIGIUM 22.222. c. 2... see eeceeeeeenecceereecmenees 1 0-2
OTHER ELEMENTS

PRIS NSPECIES | (106) veces cme cwwemce ew eneween seca aammseaneennns 5310-1
MMDUMEDNNSIIC CICS a 2-c 08. 2225. Secwideaniulesesseesunenaeeveneeunes 246 47-2
Wastracods (Tetradella?) (132) .........ccccsveesccsseseeeeeceee 66) 12-7
PRETONCROSSIGIES ICL) orcrscicca can wncwasienenaaeeerescenewermen dormeets 1 0-2
immocone fragments (15)... .....eeseseee cece eene reese ene 15 2-9
MPRMETe TOS erernicen es Arcee rattaevereecretras esenancccaasoneste i @2
| SEONG ~ peécecdnn 008885 SAH CBaSE EBS ABBE ABRe es eae niece neecon couse 45 8-6
- SORRGT'Y oSdeSeOos doe sbBospbacbhenee oABEL Sapp badge Haare udoesee 15 BSS)
Total 521- 100-2

PY = pedicle valves, BV = brachial valves, C = conjoined valves, I =
indeterminate valves, NO. = equivalent number of total animals. (N.B.
Numbers in brackets are total fragments of colonial or fragmented individuals
— see discussion). See discussion on p.127.

Leinster faunas. It is molluscan not brachiopod dominated,
with different brachiopod genera present. Five of the 11
genera are not known from the other localities and make up
10-8% of the fauna, of which only 13-4% comprises brachio-
pods. The fossils were excavated from muddy and tuffaceous
siltstones and fine sandstones at the back of the cottage.
|About 2-3 metres below the fossiliferous section are fine
/quartz pebble conglomerates with fragments of the underly-
ing slates.

_ The fossils are interpreted as having lived in a shallow
water but low energy environment, perhaps close to the shore
of the emergent volcanic cone of Grange Hill. Supporting
evidence includes the very close proximity of lavas inter-
preted as being subaerial, and the abundance of an ostracod
| (Tetradella?) which elsewhere is commonly indicative of
‘intertidal or shallow subtidal conditions.

| Kilbride, within the Upper Tramore Volcanic Formation,
iby contrast, represents a different setting. In the small
exposure there are contorted streaks and lenses of tuff in the
\dark siltstones, with very obscure bedding. The faunal evi-
\dence from this work and the limited field evidence supports
\Carlisle’s (1979: 551) interpretation of slumpedvolcaniclas-
‘tics, and that the lithology is a debris flow. The high diversity
jand low frequency of any one species suggests that faunas
from different areas and substrates were ‘sampled’ in the
downslope movement of the sediment mass. Although Carl-
isle (1979: 552) only listed certain genera as common and

129

others as ‘also in the fauna’, the assemblage listed in Table 11
shows significant differences, reflecting a different sample. In
the fauna are numerous (22) fragmentary plates of a cheiro-
crinid cystoid, of which 5 may be oral plates. These may
belong (C.R.C. Paul, personal communication 1987) to
Hadrocystis or Acantholepis (whose sole species A. jamesi
M’Coy, 1846 was described from Raheen, Waterford Har-
bour). The preservation of the cheirocrinids (Plate 16, figs
12-18), which are fractured across plates and not along
boundaries, also reflects the disturbed nature of this horizon.
There are also 9 plates of Echinosphaerites, some of which
are broken. Comparable examples of debris flows include
that described by Lockley (1984) from Builth, central Wales
and Kilbucho in Scotland (Clarkson et al. 1992).

Although it is reasonable to treat them as ‘in situ’ faunas

Table 11 Total assemblage collected from Kilbride.
Internal External Total
PV BVC I PV BVC I no. %

BRACHIOPODS
Acanthocrania? 1 1 0-6
Anisopleurella? =- 1 -- = 1 1 0-6
Bicuspina? 2 1 2 1-1
Bimuria sp. 27 - -- 1--- Piss ()
Chonetoidea cf. abdita Id! @ = = Y 2 - 14 78
Cremnorthis parva OP 7 tS Ss Hee = 8 4-5
dalmanellids, indet. ee eee ee Dee Me 3} 1-7
Hesperorthis sp. Se t= eda 3 17
Leptestiina oepiki 6 -- 12-- 6 3)93)
lingulid, indet. - 1 1 0-6
Nicolella cf. actoniae WM ==> § © == 10 5-6
‘Orthambonites’ sp. Sal ee eo 14 qi
Platystrophia sp. 2 ft gJoee= i lee 3 1-7
Porambonites sp. 2--- =-=-=-+ 2 1-1
Salopia? 1 1 0-6
Saukrodictya sp. i--- 1--- 1 0-6
Schizotreta sp. 1 1 - 1 0-6
Skenidioides costatus Ae Age ew 7 3-9)
Sowerbyella sericea 18 7. — — 138 10,— — 18 10-0
sowerbyellid, indet. 2 - 2 1-1
TRILOBITES
ey belinidicephal ong ence-cereese reenter ssasaeeer resect ners 1 0-6
Ne) nv ali(erejo) 1A oN" Sec eaeedagocermades dean sdosondadaceonansesopecoonodss 1 0-6
VAT NTONACHISI avsce ecto c seas tb = aot ee PER e eee eee eee eee 1 0-6
cybelinid librigenae (5)
Mi eerviol? 2/9) NANO . soenadadecodeedacdons sobéooanoosaocdobodnandecas 1 0-6
2GHAHOEY |DYAENUIN cooacoogscocnnooadnuacooghoscsseecoeCooasaculs 1 0-6
OTHER ELEMENTS
SONS arcoscedadaeadeddéssandeSoadsansdeonadoédasoosboscbDbDoddadoEee 1 0-6
PERSIA) OGL, _-sausseecooencdonesacseobhnoadedoooouddecobanpoapuaeopccace 6 3h)
GSI Tcf0\c Soros ance penncecunBaster caneon be sessdecansuasbnonenaccodeotcigc 3) 1-7
(NTI OLKONEY "ss sceeodonacadonneadseanatndsos oc eoosdescsosadaconc0eOsD0c00 1 0-6
ConulantidS iaedas eat tasteep eee tapes oaacasd obegieetowower tines aoe 2 1-1
PLasOporidibryOzZOaNS e.eeeeeeteresseetesese erreeeater ere eccee 16 8-9
TOSS LOA OZORINS (27) aeaceanccpscosenccadcdorannodsorcsedosbtionse 1 0-6
COVA reece once mer iscece cece aniuectee cescsleyscmtictae tc sce slieeastiress 1 0-6
CLINOLAVOSSICIESI(T7)T oecaccc sane tc serce tone tecce settee matress 1 0-6
Cheinocninidlicystoid) erases. -cereasseeeeeceeretensce eer tteer iN 9-4
cheirocrinid cystoid oral plates? (5)
JEG UI AON NUHAAHES [OENIES (6) copspdeeassococrenoononnboaecrogs.0ceC 1 0-6

Total 179 100-8

130

for stratigraphical correlation, the loose block nature of
assemblages from Ballykale (Table 12) and Clologe (Table
13) offers little information, although the fauna from Clologe
is from two distinct lithologies, one shaly and one tuffaceous.
Hiller’s (1971) collections from Frankfort and Clogh also
came from loose blocks. Carrigadaggan, although yielding a
diverse large assemblage (Table 14), gave little information
about palaeoecology, since exposure was poor and much
material came from broken blocks not completely in situ. The
coarse volcaniclastic lithology preserved details of the fossils
only poorly.

The presence of Echinosphaerites cf. granulatus M’Coy
(Plate 16. figs 1-7,9-10) in some abundance at Carrigadag-
gan, is worthy of note. It was listed as E. aurantium in the
G.S.I. Memoir (Kinahan 1879). This species is found as
almost complete individuals (6 specimens), suggesting rapid
burial as in a coarse tuff fall, but there are also numerous
isolated (15) or several associated plates (27) of the cystoid.
Forbes (1848) monographed the British and Irish cystoid
fauna, and Paul (1973, 1984) has produced a modern revision
of some diploporite and rhombiferan cystoids. This material,
which has been confirmed as Echinosphaerites (personal
communication 1987) will be dealt with in the third part of
Paul’s monograph. Bockelie (1981) has recently reviewed the
functional morphology and evolution of the genus, from
many different lithologies in Russia and Scandinavia. In spite
of the fact that there are several nearly complete thecae, the
fact that few if any preserve evidence of a stem, cover plates
of the gonopore, brachioles or oral cover plates, suggests that
they were not covered by sediment immediately after death.
It is difficult to be unequivocal about these taphonomic
factors, or the loss of periproctal plates (which occurred in
less than 5% of specimens reviewed by Bockelie, 1981: 191),
since the Carrigadaggan lithology is so coarse-grained that
fine detail of the plates is not always preserved. Two speci-
mens show projections, however, one of which is believed to
be an oral projection, the other possibly the basal plates to
which the stolon was attached (see Plate 16, figs 7, 9, 10).
Possibly the most likely explanation is that Echinosphaerites
and the cheirocrinid cystoids, of which there are 9 fragmen-
tary plates similar to the Kilbride material (Plate 16, figs
8,11), were buried rapidly by volcaniclastic material but were
subject to current action in shallow waters rather than
inundated by distal tuff falls.

Ballygarvan Bridge (Table 15) has volcanic rocks adjacent,
but exposure is too poor and the sample is too limited to infer
much. As previously noted, volcanic events may best account
for the differences between the new collections from Raheen
and the G.S.I. records (Baily in Kinahan 1882: 38-39). A
diverse fauna of 25 species including 9 trilobites in abundance
was recorded, whilst new collections (Table 16) are over-
whelmingly dominated by a dalmanellid (probably Howel-
lites, but remaining indeterminate because of the strong
deformation) with only a few trilobites and generally low
diversity. Tuff horizons (tens of cm thick) are common in the
new section that was exposed and almost certainly account
for apparent differences.

The Greenville assemblage (Table 17) also differs from
earlier collections. Since this was collected by digging
beneath a thick farmyard deposit it is most likely that
different horizons were sampled, but the cause of the changes
in assemblages from horizons in close succession remains
unknown. In contrast, the collections from Greenville-Moyne
(Table 18) probably came from the same horizon as that

M. A. PARKES |

Table 12 Total assemblage collected from Ballykale.

|
Internal External mE: |
PVABV CI SENS BVAG= aI %
Bimuria cf. dyfiensis 5910 —-—- 16 5 = = 59 81-08
Chonetoidea 1 - - 1 1-3)
dalmanellids, indet. 2) SS oa a eee |
Leptestiina sp. D, - 2 pe
Platystrophia - 1-- --- 1 2. 2a
Ptychoglyptus i= Ht. = 2a |
Saukrodictya sp. 1 = 1 1-3
Sowerbyella? 2D - 2 2-7}
strophomenid, indet. 1 1 1-3
Total 73, , 90ne |
OTHER ELEMENTS {

ramose bryozoans (5)
trilobite genal spines? (3)
trilobite thoracic segments (2)

PV = pedicle valves, BV = brachial valves, C = conjoined valves, | =
indeterminate valves, NO. = equivalent number of total animals. (N.B.
Numbers in brackets are total fragments of colonial or fragmented individuals
— see discussion). See discussion on p.127.

Table 13 Total assemblage collected from Clologe Upper.
|

Internal External Total
PVBVEG a RVeB aCe no. %

BRACHIOPODS
Anisopleurella? 1 1 - 1 1-4 )
dalmanellids, indet. 24 Ne= 16. Sea 7 10-0
Dolerorthis? toi == 1 - -- i 1-4
Glyptorthis - --- ---1 1 1-4
Kjaerina? - --1 - --1 1 1-4
Leptestiina sp. 12.1- ---=+=- 3 4:3 |
Nicolella sp. DOs NS = 2 2:9)
Oanduporella? 2: 2 2-9
Plaesiomys? 1 1 1-4 |
Ptychoglyptus sp. 1 1 1-4
Salopia? ---- -1-- i 1-4
Skenidioides sp. 1 --- 1 - - 1 1-4
Sowerbyella sp. 2 ? pes) |
strophomenid, indet. 1 2 1 1-4
TRILOBITES
tretaspid:cephiala: or.. s.as.ncetepeeber ees cesneacere -aereeee eee . 18 255i]
tretaspid cephalon fragments (14)
cybelinid librigenae (2)
AGMOSTIG Mee hata eee See ce 1 1-4
lichid cephalonqcsiinc.cosguaedh seteasceseee ceases eee eee: 1 1-4]
cheinunidicephalont a. ..-ee-seeeee-eeecees cass ease eee ee eee es 1 1-4
OTHER ELEMENTS |
mamose bryozoansy(4)) fice. aceececsesso cece. ose se eee Reece 1 1-4
prasopontdibnyOzoans pee seeeeeet creer seeteeee reese seeeeeeeraee 1 1-4
OMMNOCOME tee eaes sean secs <ecisne tecenne vase ceaarsniee teen eee eee 1 1-4
CHIN OIGFOssiclesi(4)eestees aac tee ae eee ce eaten eee 1 1-4
PASTLOPOGS Beak es pothesis es eee ee Lae eee 12 17-2!
Con lamin IAS. Ie a ease eee ne See sees 1 1-4
CORAL?” Hesse aac see saidiea-igshitiigdysemmeiandas tube gence aero 1 1-4
Onan niVGl GION Cogscsescece condeteadsondsecceconscccecuccane 3 4-3
Echinosphaerites plates (4)
LTH MOSD NCA ATAD (WN NECEY vo cos anecaodaddqebsnoseadcdsdaeaosoNaNse 1 1-4
cystoid, indet.

Total 70 96-8;

to
Ws)
— ee Et

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

Table 14 Total assemblage collected from Carrigadaggan.

131

Table 15 Total assemblage collected from Ballygarvan Bridge.

Internal External Total Internal External Total
PAY IW (CM PY EY © Il We, % PWV IEW I AY BWC TI no. %
BRACHIOPODS BRACHIOPODS
Bicuspina? cael ee 1 0:3 dalmanellid, indet. 41- - Sal = 5 6:75
Bimuria? a 1 0-3 Lingulella ovata 3 pez 2:7
Cremnorthis parva Say sb le Shel li, pate 6 1-6 Nicolella cf. actoniae 1 - 1 1 1-35
dalmanellids, indet. AMARA 25) eli eile -2 Platystrophia sp. 1i1--=- 1 Sl 1-35
Dolerorthis? 12-- -- -- 2 0-5 Salopia? a Li e=p Se al 1-35
Glyptorthis? SS Sl 1 0-3 Skenidioides sp. 1 1 | 1-35
Kullervo aff. hibernica Sea = = 31 = 4 5 1-3 Sowerbyella sp. =a = Sy SS PS 1 1-35
Leptellina cf. llandeiloensis 4 3 -— -~ -— —~ — — 4 1:0
Leptestiina oepikiampla 29 10 - - Ae Seeds YRILOBITES:
lingulid, indet. os =A 2 0:5 CEI TDEUG! DYFAIGHUTEN, coconovnccqnaancosnoncceaanooneondcannseass I 135
Nicolella cf. actoniae = > cee peat ee 7 1-8 Deacy beleicephalonwressserceseeeeee seca eee eee eee 1 1-35
ee 7 SP ; i - - 24 = 11 2:8 OTHER ELEMENTS
) Ort Pet waricata ae 4 = ; me DANO POM! TAWOZOBS — coccnssprocensuonopaoanuosccasseooessancce 20 27-0
Oxoplecia? | rs aie z Lsn Ors PATNOSS DINICZOLIS (BD) novcassocooeeseseacvocoovbnpassaecooossese 1 1-35
Platystrophia sp. See Wl 11- 9 D3 gastropods 37 50:0
Be eeriacdanjindet, = = == 2.- - 5 0:5 Eason vag Clee Rea din cing aniae's age seem dauasioete eh ees musta mee Thisatsipee
| Salopia? a x 1 0:3 one i vere: cut asbaccondsastrocdbesspstebeseebobbesboze
Me irodictya cf. sp. A Cee Mee oe a 5 13 OMMOSHACATTAD WNECA sonecsssacsocveanoacenondoccesondncdéeont 1 135
Skenidioides costatus By ae a 3 0-8 Total 74 99-95
Sowerbyella sericea 2 = eee 4 1-0
triplesiid indet. 1 --1- +--+ -- 1 0-3
triplesiid indet. 2 2) ee Oe 2 0-5 PV = pedicle valves, BV = brachial valves, C = conjoined valves, I =
indeterminate valves, NO. = equivalent number of total animals. (N.B.
TRILOBITES Numbers in brackets are total fragments of colonial or fragmented individuals
trinucleid pygidia (1) — see discussion). See discussion on p.127.
PPC LERGKCE PNA ay oct cuc sls sesesct scene sciiicscnssmeceoe messes 5 1-3
Atractopyge Peidia BA ce OS A Ie LR ee SAAR 18 4-6 Table 16 Total assemblage collected from Raheen.
| Atractopyge cephala (9)
ea (15) 5 0-5 Internal External Total
oa stcseeenenucvtvecnson it OS ES a AC hei OOS
PRPIGUIGX ICEPNAION a scccsterescseseescaserssesecesaceecdeuees 1 0-3
OTHER ELEMENTS Cen ral z ; a
RPSONOIC DTV OZOANS) .<.ceececa-aececqecceceanesaecerceeteaneense 75 19-4 ee eeeeetein © al
BeTS CHT OZ OAS) fa. eae es. Soe Sysco oautiamanens seateedse 42 10-8 pomere eves”) Oe ae ale Bh 820
PESIROPOGISPECIES, .2o0..2.0.eeeeeaeneeeenecernatsesooevenesemenece 29, «7-5 nats ae ad Lape A > 5, GunZ ; oe
>“ IEAGIIGIS - coc eqicenbsouoldadacteceentacenee ceed te eee ener aeeeeee 2 10 I ee Cable sab anthas a Sen S op Ee
DiVAIVeS ......... 2s. eeeeee cece este testes e teeter te tteetetteees 2 0-5 TRILOBITES
= CRSICIES (AO) ceansoccadesdooseesesteeemnpaeaacobppacnaecrcs 1 ae ATIDYE PYCICA atte ek hte eene tera eee 2 1-5
ort peones te ee ee eLel ele dessins dRisica vieivslieinicininic vnrialen 11 2:8 firelaspistcephalonier pete eee 1 0-7
coral’ eaiisi wees eegeearseeseeseecacsuneneeeeeeaeeaeeteneenreereeees 1 0-3 iretaspidice phialond see Cee See. SO. inane 1 0-7
Echinosphaerites single plates (15) tretaspid cephalic fragments (2)
Echinosphaerites attached plates) (27) iwressereeeeseeec eee: 5 1-3 tretaspid pygidium (1)
PIRMIMOSPNGCrIESANECAE 22.2... -cccecencecrseeeseseeneesenecoees 5 ils3}
MMMM AIC YSLOIG Meee cece ocjs sere ceasicch Ek ceeseer cee seeeceemenese 9 23 OTHER ELEMENTS
(7g 1010) (0S (3) i chase csen ana SA cea aac aba aadddcdocnclsasneaspanannncbb 1 0-7
Total 387-1001 seastiopodsisn’ weenie: ase een meee 36-265
GORE ISCO) Scpesner popradenctocdocObecods aac coohpnachaccdadosocuspadayce 2 IES)
56 2 :
PE clealves BY = brachial valves, C = conjoined valves, 1 pais BSS PREC oder CCD aC ceRRERRCt ore aecoreeceecodo tec nrorceacd g fe
indeterminate valves, NO. = equivalent number of total animals. (N.B. Me OS ie BEDE ARS AINE E OE ES LS a ¥
Numbers in brackets are total fragments of colonial or fragmented individuals Total 136 100-0

—see discussion). See discussion on p.127.

recorded by Brenchley et al. (1977) and slightly increase the
genera known from here.

Ballinatray (Table 19) preserves a rare occurrence of shelly
fossils in a normally graptolitic facies. Coastal sections of the
Ballinatray Formation north of Courtown have hitherto
“yielded only graptolites (Crimes & Crossley 1968, Brenchley
\% Treagus 1970). Shells are found in thin, densely packed
| bands in calcareous slates. Restricted exposure in the bed and
banks of the Owenavorragh River and strong deformation
make assessment difficult, but the lack of size variation,
restricted diversity and dense packing suggest that the fauna

was a coquina. Whether long distance transport or local
winnowing was involved is not clear, but all specimens are
small including the rare genera, elsewhere of a normaily
larger size.

132

Table 17 Total assemblage collected from Greenville.

M. A. PARKES

Table 18 Total assemblage collected from Greenville-Moyne.

Internal External Total Internal External Total
JAY JERAV(C Il JAY AYE I no. % PY BAY EL JAY BY (CII no. %
BRACHIOPODS BRACHIOPODS
Chonetoidea abdita ES sy = i Bil 4 = D7 Il atlas Anisopleurella sp. 8 2-- 25 -- 8 6:0
dalmanellids, indet. - --- 41- - 4 Dei], Cremnorthis parva 8 2 8 6-0
Oanduporella? le =e= 9 i= = 3 2-0 dalmanellids, indet. 39 I © = IO i = = 45 = 33g
Dolerorthis? bh 1 St - --- ly 0%
TRILOBITES Kullervo aff. hibernica 3 -- - 3 = 2) 4 3:0
RATA NAUMAN Jos CIOVBVE) soonsaocesosonccseacenooacoscaovsoace 5 3-4 Leptestiina oepiki a Or = y Oe Bu D0)
A ORAQHGOD SO. |OKBMUW soonneescnecnass0sc0nceontenaenconucs 1 0-7 lingulid, indet. 3 11 =0-748
Ampyx austinti cephala (8) Nicolella cf. actoniae Bee je A = 8 6-0
Ampyx austinii glabellar spines (7) ‘Orthambonites’? 5 ae i lke Siena
ANAM ONEE CUISUI OH PONAGVE 5 ococcaccocsunacdacosonaccspon0qquosoGnn00 16 =10-9 Onthinoerania? | 1 - 0-78
Wirinoduswagnostiformisipy eld lalaueerereeeeeceeee raat eee eee 8 34; plectambonitacean, indet. 1 0-78
Miretaspis Cephalon) .aacshiseacsnis: ashe seetmanesemaseeet cere cate 1 0-7 Rhactorthis? 1 af 1 1 (0
thinucleidicephalatee. cesceneassecercer esters eect reece Rene cereee 4 2-7 Salopia sp. S432) 2h eee
trinucleid pygidia @) Saukrodictya cf. sp. A ---- ---2 1 “07g
trinucleid thoraces (2) Skenidioides costatus fet 2S. 9 hee 1 0-75
trinucleid genal spines (2) Sowerbyella? Bh Lees Deis me 340020)
OTHER ELEMENTS strophomenid, indet. 1 = 1 0-75
ENOTOGTING! GVA! sacoccassnascasanassacooosconccesacnadoocuaeN 1 0-7 TRILOBITES
gastropods wid au Gahoiad a Mepeaame cio ueensen ane oteecastnepeetoes meter 38 ©25:9 cybelinid? librigenae (8)
bivalve _begoonoeoana9 noe sandansoconnauTaaDdooObODsUsHDOoROOAECCOLOIORC 1 0-7 indet. pygidia (1)
(OWE EIGN: Se acsascosasaedsdocaapcacode race sbecucadascnososaaaoosoas 4 2-7 indet. cephala (2) Sn ae eee 2 1-5
Total 147 100-0 OTHER ELEMENTS
erinoidiossicles: (14) steers fees eee 1 0-75
i ans 22 :
PV = pedicle valves, BV = brachial valves, C = conjoined valves, I = Bee BEYOZOADS 3-25-6200 052s soreness aaa a 4 x
indeterminate valves, NO. = equivalent number of total animals. (N.B. OSS WW OZ0ans, So aa ec oe aa aaa aia sare us OOO O5D ‘
Numbers in brackets are total fragments of colonial or fragmented individuals Cheirocrimid Cystoids ©.-....-.r.... one. eee nne eee dene een gee 3 22
—see discussion). See discussion on p.127. (OLY H (C60) 0 LS ate aaemaenoe oaoepeetnton santadbododdaserchaddiedchsonnas8 3 2:2
Total 134 100-0

Comparisons and contrasts with coeval
associations

A prerequisite to recognition of faunal provinciality in bra-
chiopod assemblages is the determination of whether appar-
ently different faunas are simply facies controlled. This was
undertaken as an integral part of this project and is briefly
summarised here. Due to the problems noted before, only
some localities yielded assemblages which could be treated as
palaeocommunities. These were Grange Hill Horizon 1,
Grange Hill House Cottage, Carrigadaggan, Raheen, Green-
ville and Greenville-Moyne.

In fact, the assemblages are clearly dissimilar to platform
sequence faunas from Laurentia, and to the marginal Scoto-
Appalachian faunas from Ireland and Britain. Few assem-
blages of similar age from the Lake District are known and
there are no modern quantitative studies for direct compari-
son. Essentially, only comparisons with Anglo-Welsh associa-
tions were applicable. Principal Component Analysis (PCA)
using Palstat (Harper & Ryan 1987) is an effective way of
discriminating and illustrating the elements which contribute
most to the variation between samples. One example is given
to illustrate the comparitive analysis undertaken. Figure 15
shows a plot of vector 1 against vector 3 for the Grange Hill
Horizon 1 association and 11 samples of the Nicolella Asso-
ciation from the Gelli-grin Formation from Lockley (1980).
Plots of vector 1 against 2 discriminated abundance trends of
dalmanellids. The plot in Fig. 15 clearly differentiated the
Grange Hill elements as contributing substantially to the
variation on vector 3. Whilst not a precise test, it shows
important differences which do not permit the association to
be considered as an equivalent to the Nicolella Association,

Table 19 Total assemblage collected from Ballinatray.

Internal External Total
PVABWVAC SIE VABWVaGaal no. %

BRACHIOPODS |
Chonetoidea sp. Lj = = Sy = l 1-8
dalmanellids, indet. 38 26 -- 810 - 14 45 78-9
Dolerorthis? 12-- --- = 2 33
Leptestiina sp. 11-- --- = i 18
plectambonitacean, indet. 1 2-- - —- — 3 4 7-0
Sowerbyella? 1 --- - -- - 1 i
TRILOBITES
TPO MT OWES? FNFBCWEL -onceadaad soaosvabesnaeses08se8eG0 2 7300802 2 3-3
OTHERS
CHINOIG OSSICIO Ss: pcieowsasatnede hess caboasete <a seenee aeepeee ens 1 18

Total 57 100-1

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

| sensu Lockley (1980). Principal of these are the abundance of
the calcareous inarticulates O. divaricata and P. harperi, and
the much greater frequency of Skenidioides and Cremnorthis
at Kildare than in the Welsh Longvillian.

Pickerill & Brenchley (1979) studied the faunal communi-
ties of the south Berwyn Hills of N. Wales, and defined
communities very similar to those of Lockley (1980). Direct
comparison using PCA was not possible due to their data
_presentation. For their Nicolella Community simple inspec-
tion shows that the Grange Hill association does not equate
(Table 20). This aspect of the study highlighted the benefits
of recording raw palaeontological data, as suggested by
Lockley (1980: 167). As Jaannusson (1984: 127) noted, there
are many published studies which cannot be compared
because of differences in tabulation of quantitative data. The
example of the Grange Hill Horizon 1 assemblage compared
with the Welsh Nicolella Association is a typical one. The
‘analysis of the suitable assemblages from Leinster in relation
to Welsh and English Caradoc (Hurst 1979a, 1979b, Pickerill
& Brenchley 1979, Lockley 1980) and especially Lockley’s
(1983) rigorous definition of eight palaeocommunities is
‘discussed in more detail in Parkes (1990). The moderate to
severe differences evident precluded placement of the Lein-
\ster assemblages into those palaeocommunities.

The localities from Leinster are in volcano-sedimentary
‘sequences associated with island arc volcanism. The rapid
‘facies changes over short distances, with little or no shelf
development and invasion of volcanic detritus into clastic
‘substrates are sufficient to explain this lack of comparable

|
|

3.9
v Skenidiodes
2.7.
Orthisocrania
z Petrocrania
15 Cremnorthis
/ v Platystrophia
F 3 vy gastropods
byell.
| a3-| ¥ ESOWer yella

1.1 0.7 2.5 4.3 6.1 7.9
V-1

Fig. 15 Principal Component Analysis of Kildare Grange Hill

| Horizon 1 association with Gelli-grin Formation Nicolella
Association samples: plot of vector 1 against vector 3. Data

| Sources: Association 1: Table 8 (p. 128), Grange Hill Horizon 1.
| Associations 2-11, Gelli-Grin Formation. From Lockley, 1980:

| fig. 12 (GGih, GGlg1, GG1b); fig. 13 (GG2d, GG2b); fig. 14

| (TB 16, TB 12); fig. 15 (R34, R30, R28).

|
|

133

Table 20 Nucleus of Nicolella Community as defined by Pickerill &
Brenchley (1979), compared to nucleus of Grange Hill Horizon |
association.

Nicolella Community Kildare, Grange Hill, Horizon |

Nicolella 34% Skenidioides 18%
Dolerorthis 21% Orthisocrania 14%
Platystrophia 16% Petrocrania 14%
Skenidioides 9% Cremnorthis 13%
Leptestiina 6% Platystrophia 9%
Eoplectodonta 4% gastropods 8%
Howellites 3% Sowerbyella 5%
Sowerbyella 3% tentaculitids 4%
Cremnorthis <1% dalmanellids, indet. 4%

community development. Lockley (1983) concluded that the
Welsh Basin, with low facies gradients for much of the
Ordovician rather than basin margin localities, provided
more stable sites for the evolution of diverse, well established
palaeocommunities.

Biogeography

The component brachiopod species of the Duncannon Group
assemblages are best understood when viewed in terms of
species migrations and palaeogeographical changes. Harper
and Parkes (1989), Harper et al. (1991) and Murphy eg al.
(1991) have dealt partially with the faunas described here. A
more complete treatment of their biogeographical context is
contained in Parkes (1992) and is not reiterated here.

In summary, these faunas are ascribed to a distinct Anglo
Welsh Province centred on Eastern Avalonia in the Caradoc.
Despite the lack of comparable palaeocommunity develop-
ment, there are numerous congeneric and conspecific forms,
many with an earlier origin in migrations from the Baltic
Province. Specific appearances of Scoto-Appalachian e.g.
Leptellina and Bimuria, and Mediterranean Province genera
e.g., Saukrodictya, can also be tracked as provinciality was
diminished by northward movement of Avalonia towards
Laurentia and low latitudes (Fig. 16, p.146). They are also
significant in constraining volcanic and tectonic events. Lavas
at Grange Hill, Kildare are tightly constrained to the Soud-
leyan — Longvillian interval. Although the stratigraphical
position of the other faunas is not so clear, the data from this
study support recent ideas that subduction-related volcanism
ceased in the mid Caradoc (Pickering et al. 1988, Leat &
Thorpe 1989). The biogeographical analysis of Ordovician
faunas is an important facet of understanding the develop-
ment of the Caledonides by Iapetus closure. Discovery of
new assemblages may provide new evidence, but revision of
faunas from old localities such as this work and Parkes (1992)
is equally important. Furthermore, new methods of analysis
such as seriation (Parkes ef al. 1990) may also reveal clearer
distribution patterns.

SYSTEMATIC PALAEONTOLOGY

For the better preserved, larger samples, the ‘material’
section of each description includes only the measured
sample on which the statistical data are based and excludes

M. A. PARKES |

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

the fragmentary material included in the Tables 8-19. For
'some genera the available material did not merit a full
| taxonomic statistical description. In these cases the best
| representative forms are figured, and for this material dimen-
| sions are given in the systematic section, if the magnification
is not an adequate indication. Full data sets are given for any
| statistically described species in a supplementary document
lodged at The Natural History Museum (see p.118) or
‘available from the author. In addition to the material
described or noted in this section, there were some poorly
preserved or fragmentary specimens, often of questionable
identity, which were not worthy of discussion but are listed in
the locality Tables (8-19). Inevitably, for each locality there
were a few unidentified specimens.

In this study, the dalmanelloid brachiopods were a com-

mon, albeit minor, element of most faunas. Although species
lineages are stratigraphically useful (e.g. Hurst 1979a), the
recognition of different genera and species is dependent on
\the measurement of small morphological variations in the
|cardinalia, musculature and shell outline, as well as aspects of
the costellate ornamentation. In the localities sampled by me,
|dalmanelloids were present in most cases, although as minor
constituents of the assemblage, except at Raheen where one
'species was dominant. Their poor preservation, small sample
numbers and frequent tectonic deformation made it difficult
|to make confident identifications; they are therefore listed as
‘indeterminate dalmanellids’ in the relevant tables. The iden-
tification of the less common elements in the faunas, such as
bivalves, ostracods, gastropods and crinoids was outside the
scope of the project, and in any case most material was poorly
preserved and generic identification would have been some-
what equivocal. However, a few elements such as the cystoid
Echinosphaerites are figured.

All material, other than that figured, is housed in the
James Mitchell Museum (JMM) in the Department of Geol-
ogy, University College Galway. The systematic classification
jof the Brachiopoda is based largely on Cocks (1978), and
Cocks & Rong (1989) for the Plectambonitoidea.

Phylum BRACHIOPODA

Class INARTICULATA Huxley, 1869

Order LINGULIDA Waagen, 1885
Superfamily OBOLOIDEA King, 1846
Family OBOLIDAE King, 1846
Subfamily LINGULELLINAE Schuchert, 1893
Genus LINGULELLA Salter, 1866
|
Lingulella ovata M’Coy, 1846 Pl. 1, figs 1-9, 11, 14

1846 Lingula ovata M’Coy: 24; pl. 3, fig. 1.

1866 Lingula ovata M’Coy; Davidson: 38; pl. 2, figs
19-23.

1866 Lingula obtusa Hall?; Davidson: 52; pl. 3, figs 31,
32.

PLATE 1

135

cf. 1963 Lingulella cf. ovata (M’Coy); Williams: 344; pl. 1,
figs 1-3.
1978 Lingulella ovata M’Coy; Cocks: 15, 171, 172.
cf. 1980 Lingulella cf. ovata (M’Coy); Lockley: 203, fig. 25.
cf. 1980 Lingulella sp. A; Hiller: 123, figs 11-15.

MATERIAL AND LOCALITY. Kildare, Grange Hil!, Horizon 2:
8 single valves, 1 conjoined pair of valves.

DESCRIPTION. Exterior. Very large equally biconvex, elon-
gately oval valves with acute beak. Maximum wicth at 66% of
length although sides nearly parallel. Shell variably buff,
brown and black coloured, ornamented with fine concentric
growth lines on lateral and anterior margins, nearly smooth
centrally. Occasional faint radial striations medially towards
anterior margin. Depth of valves 10% of length. Width is
60% of length.

Ventral interior. Not positively known, but one specimen
shows smooth surface with faint median striations towards
anterior margin.

Dorsal interior. Two specimens show dorsal median sep-
tum, extending 50% and 70% of the length, «expanding in
width and height anteriorly. The growth lines ere deflected
anteriorly by the median septum.

DiscussION. M’Coy (1846) described the species from three
localities; Ballygarvan Bridge, Kildare and Newtown Head,
Co. Waterford. Cocks (1978) selected the lectotype
(NMING:F4578) from the Chair of Kildare. He stated it was
from the Kildare Limestone Formation (Rawtheyan), but
M’Coy recorded it as ‘common in the shale of Chair of
Kildare’ which is actually the lithology of the Grange Hill
locality. The opportunity is taken here to figure some good
topotype material. The Griffith Collection specimen in the
N.M.I. (F5482), a paralectotype from Ballygarvan Bridge,
accords with the type material in its proportions. although it
is only the anterior half of a valve. A significant feature
revealed by a larger sample is the very large sive which this
species can attain. M’Coy (1846) records the length as one
inch three lines (c. 32 mm) but the largest specimen collected
is 43 mm long, although much smaller specimens are also
present. It appears that this large size is probably normal for
the species rather than representing a population which
realised optimal conditions for growth, as suggested by
Pickerill (1973) in a study of Lingulasma tenuigranulata from
North Wales. Harper (1984: 19), in noting the similarity of L.
carrickensis carrickensis to L. ovata, raised the problems of
Ordovician Lingulella taxonomy. The sample described here
may help facilitate a future multivariate morphometric analy-
sis of described species. Moreover L. ovata may prove
ultimately to be a synonym of L. brevis (Portlock, 1843), as
noted by Mitchell (1977). The relative proportions are similar
but the Pomeroy sample is considerably smaller in mean size.

igs 1-9, 11, 14 Lingulella ovata M’Coy. Kildare, Grange Hill Horizon 2. 1, BC 12634a, interior, x 2. 2, BC 12369, exterior, x 2. 3, BC
12635a, exterior, x 13. 4, BC 12636, exterior, x 3.5, BC 12637, exterior, x 2. 6, BC 12638, interior, x 137. 7, BC 12634b, interior, x 2.
8, BC 12640b, interior, x 2. 9, 11, 14, BC 12640a, internal mould, counterpart of Fig. 8; 9, ventral view, x 2; 11, dorsal view, x 14;

enlarged dorsal view showing median septum, x 2.

ig. 12 Schizotreta cf. corrugata Cooper. Kilbride. BC 12641b, pedicle valve exterior, x 4.
Figs 10, 13 Acanthocrania? sp. Kildare, Grange Hill Horizon 1. Brachial valve. 10, BC 12642a, internal mould, x 10. 13, BC 12642b, external

mould, x 10.

Superfamily DISCINOIDEA Gray, 1840
Family DISCINIDAE Gray, 1840
Subfamily ORBICULOIDEINAE Schuchert & Le Vene,
1929
Genus SCHIZOTRETA Kutorga, 1848

Pl. 1, fig. 12

1956 Schizotreta corrugata Cooper: 277; pl. 21, figs 7-20;
pl. 28, figs 24-28.
cf. 1977 Schizotreta cf. corrugata Cooper; Mitchell: 25; pl.
1, figs 26-30.
cf. 1984 Schizotreta cf. corrugata Cooper; Harper: 39; pl. 4,
figs 3-5.

Schizotreta cf. corrugata Cooper, 1956

MATERIAL AND LOCALITY. Kilbride: a single internal and
external mould of a pedicle valve.

DESCRIPTION. Almost circular, transversely elliptical shell,
with length 95% of width. Ornament of raised concentric
ridges crowded together on either side of the umbo, number-
ing 6 per mm between 3 and 4 mm anterior of the approxi-
mate position of the umbo. Details of umbo not seen owing to
poor preservation.

Ventral interior. Smooth, preservation too poor for detail
except faint limbus. (Dorsal valve unknown.)

MEASUREMENTS. BC 12641: (X1) length = 76 mm, (X2)
width = 8 mm (PI. 1, fig. 12).

DISCUSSION. The size, shape and ornament suggest this is
very similar to S. corrugata, especially in the asymmetry of
the apex and the lateral profile of slightly concave posterior
slope and flat anterior slope. The species is recorded from
Pomeroy and Girvan although in similarly sparse numbers.

Suborder CRANIIDINA Waagen, 1885
Superfamily CRANIOIDEA Menke, 1828
Family CRANIIDAE Menke, 1828
Genus ACANTHOCRANIA Williams, 1943

Piste ties 10513

MATERIAL AND LOCALITY. Kildare, Grange Hill, Horizon 1:
A single internal and external mould of a brachial valve.

MEASUREMENTS. BC 12642b: X1 = 4-6 mm, X2 = 5-0 mm,
X4 = 2-3 mm (PI. 1, fig. 13).

Acanthocrania? sp.

DIsCUsSsION. The preservation of this single valve, equivo-
cally assigned to Acanthocrania, is poor. It is steeply conical
with umbo? closer to the posterior margin. The depth is 50%
of the length, a figure close to the Portrane species A.
cracentis Wright, 1963, but greater than the Acanthocrania.

PLATE 2

M. A. PARKES |

|
sp. of Mitchell (1977) from the Killey Bridge Formation at |
Pomeroy. The internal features are not seen, but the exterior |
has an impression of an oblique ribbing pattern.

Genus ORTHISOCRANIA Rowell, 1963

Orthisocrania divaricata (M’Coy, 1851) _ PI. 2, figs 1-8 !

1846 Crania antiquissima ? Eichwald; M’Coy: 25.

1851 Pseudocrania divaricata M’Coy: 187; pl. 1, H, figs 1, _
Dy.

1852 Pseudocrania divaricata M’Coy, in Sedgwick &
M’Coy: pl. 1, H, figs 1, 2.

1853 Crania divaricata Davidson: 122; pl. 4, figs 246, 247.

1858 Crania catenulata (Salter MS); Baily, in Jukes et al.: |
@). Tah. 3).

1859 Crania divaricata Salter: 212, fig. 2.

1866 Crania (Pseudocrania) divaricata M’Coy; Davidson: —
78; pl. 8, figs 7-12. |

1875 Crania divaricata M’Coy; Baily: 32; pl. 11, fig. 5. |

1963 Orthisocrania divaricata M’Coy; Rowell: 39. |

cf. 1963

Pseudocrania cf. divaricata M’Coy; Williams: 345; |
pl. iefiesi4s: .
1965 Orthisocrania divaricata M’Coy; Rowell, in Will- |
iams et al.: H290, fig. 181, 6a—6c. |

1970 Orthisocrania divaricata M’Coy; Wright: 97.
? 1976 Orthisocrania sp.; Neuman: 19; pl. 1, figs 1, 2.

1978 Orthisocrania divaricata M’Coy; Cocks: 30.
? 1980 Orthisocrania sp.; Lockley: 205, figs 31a, b. |

MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon
1: 41 external moulds, 10 internal moulds of brachial valves,
14 internal moulds of pedicle valves, 2 indet. internal moulds. |
Kildare, Grange Hill, Horizon 2: 1 internal mould of a
pedicle valve, 3 internal moulds of brachial valves, 2 external |
moulds. Carrigadaggan: | internal mould of a pedicle valve, 1
internal mould of a brachial valve.

DESCRIPTION. Exterior. Unattached, shallowly biconvex | —
equidimensional valves. Outline is subcircular to subquadrate
with mean length equal to width, although mean position of
maximum width (X3) is at 62% of the length from the |
posterior margin. Shell is calcareous and punctate. Ornament
of fine costellae, mean value of 15 per 5 mm at 10 mm to the
anterior of the beak. Growth is mixoperipheral from mar-
ginal beak, occasionally holoperipheral in larger specimens.
Faint concentric growth lines on outer margins of larger
specimens.

Ventral interior. The broad flat limbus is widest at the
anterior and posterior shoulders, non-pustulose and is
approximately 30% of the width. Large oval or elliptical
anterior adductor scars are slightly elevated from shell floor. |

Figs 1-8 Orthisocrania divaricata (M’Coy). Kildare, Grange Hill Horizon 1. 1, BC 12643, latex cast of external mould, x 4. 2, BC 12644,
external mould, x 4. 3, BC 12645, internal mould of brachial valve, x 2. 4, BC 12646, internal mould of pedicle valve, x 2. 5, BC 12647,
internal mould of brachial valve, x 2. 6, BC 12648, latex cast of internal mould of brachial valve, x 2. 7, BC 12649, internal mould of

pedicle valve, x 2. 8, BC 12650, internal mould of pedicle valve, x 2.

Figs 9-15 Petrocrania harperi sp. nov. Kildare, Grange Hill Horizon 1. 9, 12, BC 12651, external mould of brachial valve, and latex cast, both
x 4. 10, BC 12652. internal mould of brachial valve, x 4. 11, Holotype BC 12653, internal mould of brachial valve, x 4. 13, BC 12654,
internal mould of brachial valve, x 4. 14, BC 12655, internal mould of brachial valve, x 4. 15, BC 12656, latex cast of external mould of

brachial valve, x 4.

Figs 16-17 Philhedra sp. Kildare, Grange Hill Horizon 1. BC 12657, latex cast of external mould of brachial valve, lateral and dorsal views, X

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

sh wow. p
Rah Ped K

138

The posterior scars are smaller and less impressed. The
anterior half of the shell is marked by numerous narrow
ridges with broader flat interspaces, giving a very distinct
pallial sinus pattern. The anterior scars are divergent from
the posterior side of the mid length, towards the postero-
lateral margins.

Dorsal interior. Similar in most respects to the ventral
interior. Additional muscle scars on a small raised platform
sagittally anterior of the anterior adductor scars.

MEASUREMENTS
External moulds:

Variates Xl x2 x3
Means 16-4 16:8 9-79

Sample size 40 4] 40
Variance-covariance 14-56 8-33 8:34
Matrix 15-0 4-38
6.20

Brachial valve internal moulds:

Variates YAN DOD 3} OS) OD)
Means 16-6) 17-2) 10-0 Re2- 668-23
Sample size 10 10 10 10 9

Variance-covariance Cay 3A] Set? (93) 116 3)7/

Matrix 6:98 4:19 0-97 —0-96
5:76 0-91 1-06
0-28 0-03
1-97
Pedicle valve internal moulds:
Variates Xl x2 x3 X8 X9
Means 17-9 1-6) Wit 2:99 8-95
Sample size 13 14 12 14 11
Variance-covariance 8-01 0-60 3-78 0:28 1-79
Matrix 9-40 0:39 0-71 0-08
PT Oris) jIk@s)
0-19 —0.11
1.62

DiscussION. Davidson (1866) provided the most complete
description of this species. Although Rowell (1963) diag-
nosed the genus his description is sparse and no specimens
are figured. The sample described herein is the first modern
description of a reasonably large sample of the species with
detailed morphological measurements. Rowell (1963) gave a
detailed history of the complex taxonomic types, and clearly
differentiated between Orthisocrania and the similar
Pseudocrania on the basis of external ornament and pseudo-
interareas present in the former genus. Other modern sys-
tematic records, e.g. Williams (1963), Lockley (1980) and
Neuman (1976), have described only very limited or dubious
material, although the preservation of Williams’ (1963: 345,
fig. 4; pl. 1, fig. 5) brachial valve internal mould enabled him
to interpret the interior impressions better than the material
described herein. This material is too inadequately preserved
to be unequivocal about the presence of pseudointerareas.

In the absence of modern taxonomic references to the

M. A. PARKES |}

Baltic species of the genus, the principal characters separat- | |
ing them remain unknown. Published figures (Heune 1899) |)
and photographs (from D.A.T. Harper) of Baltic specimens
show little difference, although the mean number of costellae
in O. divaricata is apparently less than in O. planissima.
Modern revision may show many of these species to be |
synonymous.

Genus PETROCRANIA Raymond, 1911

Petrocrania harperi sp. nov. Pl. 2, figs 9-15

NAME. For Dr D.A.T. Harper.

DIAGNOsIs. Variably convex to conical brachial valve of oval
to subcircular outline, wider than long. Posterior slope to
apex concave, anterior slope convex. Apex situated at one
third length from posterior margin; ornament of concentric |}.
growth lines, coarser peripherally. Dorsal interior with char- |.
acteristic pair of circular anterior muscle scars between apex |}
and mid length. Posterior scars not seen or poorly impressed. ||.
Narrow limbus sometimes developed. Ventral valve
unknown.

HOLOTYPE. BC 12653 (PI. 2, fig. 11). Paratypes BC 12651-2,
BC 12654-6. Kildare Grange Hill Horizon 1.

MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon
1: 32 internal moulds of brachial valves, 4 external moulds of
brachial valves. Kildare, Grange Hill, Horizon 2: 2 internal
and 4 external moulds of brachial valves.

DESCRIPTION. Exterior. Shallowly convex to conical brachial
valves with apex situated posteriorly. Shell outline generally
oval but variable from subcircular to occasionally subtriangu-
lar. Length less than width, 90% mean value, and mean
depth 45% of length. Apex situated at mean value of 36% of
length. Maximum width at 60% of length. Posterior slope to
apex concave, occasionally flat or convex. In a few specimens
the apex overhangs posterior margin. Anterior slope from
apex convex, occasionally flat. Anterior profile conical to) |
evenly convex. Ornament of concentric growth lines, variable | |
from fine to coarse, generally coarser peripherally. Pedicle
valve unknown.

Dorsal interior. Characteristic pair of circular anterior
muscle scars as faint raised areas, situated at position
between apex and 51% of length of shell. Smooth interior | |
with faint posterior scars occasionally seen. Some moulds} |”
have a faint narrow limbus. (Ventral interior unknown).

MEASUREMENTS
Brachial valve internal moulds:

Variates Xl, X2 X38 X40x9) Xie
Mean 8:57 9-88 5:08 3-88 5-04 3-029
Sample size 32 32 BR) BN 30

Variance-covariance 5-92 6:26 3-88 2-35 1-01 1-93 |,

Matrix 10-46 3:54 3:21 1-17 1-89
3-09 1-58 0:37 4-32

2:42 0:0 1-02

1:10 0-48

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

DiscuSSION. In spite of the variability in a moderately large
“sample the species is well defined, although it is only doubt-
‘fully assigned to Petrocrania. This species does not possess
the radial ornament characteristic of Philhedra (Wright
1963), neither does it have the spinose ornament of Acan-
thocrania, although craniid taxonomy is in need of revision.
Since the status of Philhedrella is in some doubt (Wright
1963; Harper, 1989 personal communication) this material is
assigned to Petrocrania on the basis of the external ornament.
This is in spite of its apparently larger anterior adductor scars,
supposedly a characteristic of Philhedrella (Wright 1963), and
although the reverse is considered an important feature of
Petrocrania. However, the material described here may sim-
oly have poorly impressed posterior scars, or even be rela-
tively juvenile specimens of a large species for the genus. The
nternal morphology does not show much similarity to any
described species. It resembles Petrocrania dubia Williams
1974), from the Soudleyan Whittery Shales in the Shelve
district of Shropshire, in having (probably) larger anterior
adductor scars, but differs in the apex being situated posteri-
orly rather than medially. Pedicle valves were not found in
his sample.

Genus PHILHEDRA Koken, 1889

Philhedra sp. Pl. 2, figs 16, 17

MATERIAL AND LOCALITY. Kildare, Grange Hill, Horizon 1:
i single external mould of a brachial valve.

DESCRIPTION. Exterior. Irregularly conical subcircular con-
vex valve, slightly asymmetrical apex. Ornament of irregular
adial ribs, from apex to margins of valve. (Jnterior
inknown.)

a BC 12657: (X4) depth = 9 mm, (X2) max.
liameter = 124 mm (PI. 2, figs 16, 17).

DISCUSSION. The distinctive radial ornament is sufficient to
istinguish this from the similar Petrocrania sp. found in
‘ibundance at the same locality, which has an ornament
concentric to the apex. It also separates it from Philhedrella,
vhich is very similar internally to Philhedra and Petrocrania
yut has no radial ornament. Although no interiors are
Ositively assigned to Philhedra, there is a possibility that
ome of the specimens counted as Petrocrania in the total
ounts at Kildare may in fact belong to Philhedra sp. How-
_ ver, the distinctions between the genera are complicated by
€ inadequately known type species, discussed by Wright
1963: 251). Pending a future revision of the three crantid
enera, the specimen is best assigned to Philhedra. Wright
1970) recorded neither Petrocrania nor Philhedra from Kil-
are.

139

Class ARTICULATA Huxley, 1869
Order ORTHIDA Schuchert & Cooper, 1932
Superfamily ORTHOIDEA Woodward, 1852

Family ORTHIDAE Woodward, 1852
Subfamily ORTHINAE Woodward, 1852
Genus ORTHAMBONITES Pander, 1830

‘Orthambonites’ spp. Pls3> figs 1—5

MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon
1: 6 internal moulds of brachial valves, 2 internal moulds and
2 external moulds of pedicle valves, both incomplete. Kil-
bride: 8 internal and 1 external moulds of pedicle valves, 14
internal and 3 external moulds of brachial valves.

MEASUREMENTS (mm)

X1 xe
P1.3, fig. 1. 10:5 12-6
P1.3, fig. 2 4-0 4-6
P1.3, figs 3, 4 6-6 9-0
P13, fig. 5 5-2 73

DISCUSSION. The small samples are inadequate to justify
assignment to any species and in any case the taxonomy of
‘Orthambonites’ is in need of revision.

Subfamily PRODUCTORTHINAE Schuchert & Cooper,
1931
Genus NICOLELLA Reed, 1917

Nicolella cf. actoniae (J. de C. Sowerby, 1839)
Pl. 3, figs 6-9, 11

MATERIAL AND LOCALITIES. Kilbride: 10 internal and 8
external moulds of pedicle valves, 6 external moulds of
brachial valves; all material is fragmentary. Carrigadaggan: 3
internal and 8 external moulds of pedicle valves, 1 internal
and 5 external moulds of brachial valves; material is mostly
fragmentary. Greenville-Moyne: 3 internal and 7 external
moulds of brachial valves, 3 internal and 4 external moulds of
pedicle valves and 1 conjoined internal mould. Kildare,
Grange Hill, Horizon 1: 1 internal and 1 external mould of a
brachial valve.

DISCUSSION. Williams (1974: 58) commented on the mor-
phological stability of Nicolella actoniae from mid Caradoc to
early Ashgill times. All the samples here are probably very
close to N. actoniae, or the subspecies N. actoniae obesa
Williams (1963) from Bala, North Wales, a much deeper
form than the nominate subspecies. All the samples collected
here are poorly preserved and mostly broken, as well as
relatively small in numbers; hence measurements of variation
and counts of rib numbers are not possible. Thus assignment
to one species, or several, is unrealistic until larger samples
are available to assess the variation in morphology.

The Greenville-Moyne sample is apparently not as deep in
the ventral valve as that from Carrigadaggan or Kilbride, but
this may be a preservational effect, owing to disparity in
compaction between the fine-grained mudstones _ of
Greenville-Moyne, and the coarse volcaniclastics of Carri-
gadaggan or the tuffaceous siltstones of Kilbride. The latter

M. A. PARKES |

140

| BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

two samples are strongly convex in the pedicle valve and
| appear most similar to N. actoniae obesa, from the Gelli-grin
‘Formation of the Bala district. The broken nature of the
samples, particularly of the Kilbride one (as are most species
from there), generally precluded counts of rib numbers, but
‘three brachial valve external moulds from Kilbride had 10, 12
/and 15 ribs, and one pedicle valve had 11 in total.
__ Type specimens of N. interplicata (M’Coy, 1846), from the
/Kildare Limestone Formation, were examined (NMI-
‘lectotype F4565, paralectotypes F5564 and F11604), but they
are inadequate for formal comparison. Indeed the latter
specimen is probably not a Nicolella at all. As noted by Cocks
(1978) the evaluation of topotypes from the Kildare Lime-
stone Formation is necessary to assess this species. Whether
| Nicolella calcarata M’Coy (1846), cited as rare in the slates of
‘Greenville and very common in the slates of Slieveroe,
Rathdrum, is simply a deformed version of N. cf. actoniae or
a separate species is not clear, since no Greenville topotypes
were recovered. The lectotype (F4567) is broken and did not
appear similar to any of the samples, being more transverse
‘and coarsely costate. A paralectotype (F5509) from Slieveroe
jis also deformed and broken but appears similar to N. cf.
actoniae.

Larger collections of better-preserved material would be
‘desirable to enable both a biometric comparison with N. cf.
actoniae and described subspecies as well as pre-Longvillian
species such as N. humilis (Whittington & Williams 1955) and
N. cf. strasburgensis (Williams 1962). Provisionally, the
present material is compared to N. actoniae which, as pres-
ently understood, encompasses a wide range of variation and
a lengthy time span (mid Caradoc — mid Ashgill).

\Nicolella ? sp. Pieseticele

MATERIAL AND LOCALITY. Kilbride: 1 internal and external
{mould of a brachial valve.

DiscussION. This single concave brachial valve is question-
ably assigned to Nicolella, but is different from the other
samples of the genus, including the Kilbride material, princi-
pally in being large and having a much greater ribbing
frequency (at least 26 ribs). The brachial interior has the erect
cardinal process, short divergent brachiophores and heavy
deposits of secondary shell ankylosing the brachiophores to a
short median ridge which are characteristic of Nicolella. It
may possibly be compared to N. asteroidea Reed, which has
more ribs than N. actoniae, but the preservation of the
specimen is inadequate to assess the branching pattern of
costellae.

PLATE 3

L 12661, internal mould of pedicle valve, x 4.

141

Family DOLERORTHIDAE Opik, 1934
Subfamily HESPERORTHINAE Schuchert & Cooper, 1931
Genus HESPERORTHIS Schuchert & Cooper, 1931

Hesperorthis sp. Else ie 10

MATERIAL AND LOCALITY. Kilbride: 3 internal moulds of
pedicle valves.

DIscussION. The small numbers of the genus are inadequate
for specific determination, but the long apsacline interarea
and overall shape suggest the specimens belong to Hesperor-
this. The genus is known from the Caradoc rocks of Grange-
geeth, as the Estonian species H. inostranfzefi, but the
Kilbride species is dissimilar and much smaller. Two species
are known from Girvan (Williams 1962) and one from the
Llanvirn of Wales (Lockley & Williams 1981), but formal
comparison requires a better preserved and larger sample
from Kilbride.

Family PLAESIOMIIDAE Schuchert, 1913
Subfamily PLAESIOMIINAE Schuchert, 1913
Genus PLAESIOMYS Hall & Clarke, 1892

Plaesiomys multiplicata Bancroft, 1945
; Pl. 3, figs 13-16; Pl. 4, figs 1-6; Pl. 7, fig. 12
21896 Orthis flabellulum Sowerby; Reynolds & Gardiner:
589.
1945 Dinorthis (Plaesiomys) multiplicata Bancroft: 244;
pl. 35, figs 4-6; pl. 36, figs 1-3.
1968 Dinorthis multiplicata Bancroft;
Romano: 47; pl. 5, fig. M.
1978 Plaesiomys multifida (Salter); Cocks: 50 (pars).
1978 Dinorthis multiplicata Bancroft; Brenchley: 160.
1980b Lordorthis sp.; Mitchell, in Romano: 206.
cf. 1985 Plaesiomys cf. multiplicata Bancroft; Harper et al.:
291; figs 6-24.

Diggens &

MATERIAL AND LOCALITY. Kildare, Grange Hill House Cot-
tage: 26 internal and 12 external moulds of pedicle valves; 13
internal and 20 external moulds of brachial valves.

DESCRIPTION. Exterior. Large, dorsibiconvex to convexi-
plane valves of rounded subquadrate outline, with maximum
width just anterior of hinge line, and about two-thirds as long
as wide. Evenly convex anterior and lateral profiles in
brachial valve, but pedicle valve flat to weakly concave
except for swollen posterior axial surface. Ventral interarea
flat and apsacline with open delthyrium. Dorsal interarea flat
and orthocline, with open notothyrium. Radial ornament of
evenly rounded costae and costellae and interspaces, and

Figs 1-5 ‘Orthambonites’ spp. 1, 34 Kildare, Grange Hill Horizon 1. 1, BC 12658, internal mould of brachial valve, x 4. 3, 4, BC 12660,
internal mould of brachial valve, latex cast and mould, x 4. Figs 2, 5 Kilbride. 2, BC 12659, internal mould of brachial valve, x 4. 5, BC

igs 6-9, 11 Nicolella cf. actoniae (J. de C. Sowerby). 6-7, 9 Kilbride. 6, 7, BC 12662, internal mould of pedicle valve, latex cast and mould,

x 4. 9, BC 12664a, internal mould of pedicle valve, x 2. Figs 8, 11 Greenville-Moyne. 8, BC 12663b, external mould of pedicle valve, x 2.
11, BC 12663a, internal mould of pedicle valve, counterpart of Fig. 8, x 2.
Fig. 10 Hesperorthis sp. Kilbride. BC 12666, internal mould of pedicle valve, x 10.

fe 12 Nicolella? sp. Kilbride. BC 12665b, external mould of brachial valve, x 2.

igs 13-16 Plaesiomys multiplicata Bancroft. Kildare, Grange Hill House Cottage. 13, 16, BC 12667, internal mould of brachial valve, and
| latex cast, x 4. 14, 15, BC 12668, external mould of pedicle valve, and latex cast, x 4.

f
I
|
/

M. A. PARKES

142

| BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND
very fine concentric growth lines. Counts of 18-25 costae and
'costellae are present on 2, 0, 0, 1, 1,1, 0 and 1 valve exteriors
‘at the 5 mm growth stage and counts of 27 and 31 costae and
‘costellae on 2 and 1 valve exteriors at the 10 mm growth
stage.

| Ventral interior. Stout, small teeth directed dorsilaterally
from anterior margins of wide delthyrium are supported by
‘strong receding dental plates. External ornament strongly
‘impressed, particularly near anterior margins.

Dorsal interior. Simple, linear cardinal process is slightly
‘thickened posteriorly, situated on notothyrial platform which
extends anteriorly for one-third of valve length as a low broad
ridge. Blade-like, divergent brachiophores supported by
\stout bases which, with the hinge line, define deep sockets.

|
‘Discussion. Although large, the sample cannot easily be
‘statistically compared to the existing descriptions of the
‘species because many of the specimens are broken or poorly
preserved. However, sufficient distinctive material is
\described to assign it confidently to the species illustrated by
‘Bancroft (1945) from the Soudleyan of Glyn Ceiriog, North
Wales, and in particular the sample described by Harper &
\Mitchell (Harper et al., 1985) from the Clashford House
Formation of Co. Meath. Re-collection of more specimens
will allow a better assessment of the variability in the species,
!particularly in external ornament, since the available material
|shows a few specimens more like the closely related genus
Dinorthis, than like Plaesiomys. The relationship of these two
genera is in need of reassessment. However, the record of
Orthis flabellulum from here (Reynolds & Gardiner 1896) is
probably the present species. Lamont (1953) noted that it was
‘apparently a late variety of J. de C. Sowerby’s species with
‘bifurcation and trifurcation of ribs’. The specimens collected
by Lamont are now held in the National Museum of Ireland,
but were apparently never figured or described. They are
labelled as ‘Dinorthis peplos’ on Lamont’s labels, but no
publication of this name is known.

Family PLECTORTHIDAE Schuchert & Le Vene, 1929
Subfamily PLATYSTROPHINAE Schuchert & Le Vene,
1929
Genus PLATYSTROPHIA King, 1850

Platystrophia sp. 1 Pl. 4, figs 7-17; Pl. 5, figs 1-3

MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon
1: 30 internal and 17 external moulds of pedicle valves; 33

internal and 13 external moulds of brachial valves; 6 internal

and 10 external conjoined moulds; 12 external fragments.

(Most of the material assigned to Platystrophia is incomplete.

Kildare, Grange Hill, Horizon 2: 1 conjoined shell and one
rachial valve.

143

Discussion. Although the large sample of specimens from
Grange Hill, Kildare, was clearly identifiable as Platystrophia
in both internal and external moulds, the material is almost
all fragmentary or partially broken, precluding valid mea-
surements. A statistical assessment was not possible. Since
Platystrophia is a ubiquitous genus in Middle and Upper
Ordovician rocks in Europe and America, with little variation
amongst the many described species, it is necessary to reiter-
ate the need for a complete species revision. Many authors
such as Williams (1962: 126; 1963: 371), Wright (1964: 206),
Cocks (1978: 55) and Hiller (1980: 143) have discussed the
artificial nature of Cumings’ (1903) scheme of species group-
ings, elaborated further by McEwan (1920) and modified in
terminology by Schuchert & Cooper (1932: 67). In this
scheme, the present material is all placed in the bicostate
group, with 2 costae in the ventral sulcus and 3 on the dorsal
fold. The counts on suitable material showed there were 1, 5,
3 and 2 pedicle valves with 4, 5, 6 and 7 costae respectively,
on each flank.

The Kildare population is finely pustulose, but distinguish-
ing it from other species is a concentric ornamentation
inviting comparisons with P. caelata Williams from the Soud-
leyan of Shelve, Shropshire (Williams 1974: 76-77; pl. 12, figs
13, 14, 16-19). The ornament is of differentially developed
lamellae, but further investigation would be needed to assess
whether the lamellae are of the distinctive P. caelata type or
merely accentuated growth lines. In his description Williams
stated that P. caelata is uniplicate, but later noted all speci-
mens are biplicate (bicostate); this would appear to be correct
from the figures.

Platystrophia sp. 2 PI. 5, figs 4, 5

MATERIAL AND LOCALITY. Kilbride: 1 external and 3 inter-
nal moulds of brachial valves; 1 internal and 1 external
moulds of pedicle valves, all incomplete.

DISCUSSION. The present material is inadequate for specific
determination but appears to differ from Platystrophia sp. 1
from Kildare, Grange Hill in having 7 ribs on the ventral
flanks, although being of smaller average size. It also lacks
the strongly developed overlapping lamellae, although in
other respects it is similar, belonging to the bicostate group.

Subfamily RHACTORTHINAE Williams, 1963
Genus RHACTORTHIS Williams, 1963

PI. 5, figs 6-11

MATERIAL AND LOCALITY. Kildare, Grange Hill, Horizon 1:
4 internal and 2 external moulds of brachial valves, 2 external
moulds of pedicle valves and 1 conjoined internal and exter-
nal mould.

Rhactorthis sp.

LATE 4

igs 1-6 Plaesiomys multiplicata Bancroft. Kildare, Grange Hill House Cottage. 1, 2, BC 12669, internal mould of pedicle valve, and latex
cast, x 4. 3, 4 BC 12670, external mould of brachial valve, latex cast and mould, x 2. See also PI. 7, fig. 12. 5, 6, BC 12671, internal mould

of brachial valve, latex cast and mould, x 4.

Figs 7-17 Platystrophia sp. 1. Kildare, Grange Hill Horizon 1. 7, BC 12672, internal mould of pedicle valve, x 2. 8, BC 12673, latex cast of

external mould of brachial valve, x 4. 9, BC 12674, latex cast of external mould of pedicle valve, x 4. 10, BC 12675, latex cast of external

mould of pedicle valve, x 2. See also Pl. 5, fig. 2. 11, 17, BC 12676, internal mould and latex cast of brachial valve, x 4. 12, 15, BC 12677,
external mould and latex cast of brachial valve, x 4. See also PI. 5, fig. 1. 13, BC 12678a, internal mould of pedicle valve, x 2. 14, BC
12679, latex cast of external mould of brachial valve, with a Petrocrania attached, x 2. 16, BC 12680a, internal mould of brachial valve, x

M. A. PARKES

144

) BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

MEASUREMENTS (mm)

| XX. XS

PI.5, figs 8,9 8-1 10-4 3-1 (conjoined internal
mould)

PLS, fig.10 8-2 11-1 4-1 (brachial valve)

PI.5, figs 6,11 9-8 9-5 — (strained brachial

valve)

Discussion. The sparse numbers of this genus from Kildare
are inadequate to justify assignment, either to the type
species R. crassa Williams from the Longvillian of Bala, or to
2ither of the other Caradoc species actoniae and grandis
erected by Hurst (1979a), from the type Caradoc of Shrop-
shire. Its presence, however, serves to emphasize the similari-
ties of the Kildare fauna to that of the Bala district of North
Wales.

Family CREMNORTHIDAE Williams, 1963
Subfamily CREMNORTHIDAE Williams, 1963
Genus CREMNORTHIS Williams, 1963

Cremnorthis parva Williams, 1963

| Pl. 5, figs 12-16; Pl. 6, figs 1-7
1963 Cremnorthis parva Williams: 379; pl. 4, figs 15-23;
text-fig. 9.

MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon
|: 38 internal moulds of brachial valves, 31 internal moulds of
yedicle valves; 14 external moulds of brachial valves, 7
external moulds of pedicle valves. Carrigadaggan: 3 internal
ind 1 external moulds of pedicle valves, 3 internal moulds of
yrachial valves. Kilbride: 9 internal and 1 external moulds of
pedicle valves, 6 internal moulds of brachial valves.
sreenville-Moyne: 8 internal moulds of pedicle valves and 2
nternal moulds of brachial valves.

DESCRIPTION. Exterior. Subcircular to semicular outline,
noderately biconvex, with both valves about three-tenths as
leep as long. Maximum width occurring at less than one-third
he length of the shell. Length of brachial valve about
even-tenths of the width. Pedicle valve length about four-
ifths of width. Brachial valve gently sulcate, with flatly
onvex lateral profile. Dorsal interarea short and anacline,
entral interarea apsacline and about a quarter the length of
he valve. Radial ornamentation costellate with angular cos-
ae and costellae about 5 per mm at 2 mm anterior of
imbones. Shell impunctate.

| Ventral interior. Short teeth connected to shell floor by

»LATE 5

pedicle valve (probably a juvenile specimen), < 10.

145

thickened deposits, and long apsacline interarea, which
together bound a deep umbonal cavity. Subtriangular muscle
scar does not extend much beyond cavity anteriorly. Muscle
field composed of wide median adductor scars flanked by pair
of narrow diductor scars which are lobate anteriorly. Sagittal
length of muscle field nearly two-fifths length of valve.
Dorsal interior. Large cardinal process, standing above
hinge line, is continuous with blade-like median septum
which extends four-fifths of the valve length, and is highest at
about mid-length. Brachiophores short and slightly diver-
gent, continuous with bases which curve posterolaterally as
fulcral plates to define subtriangular sockets. Brachiophore
bases are situated at about a quarter of the valve length.

MEASUREMENTS
Pedicle valve internal moulds (Kildare, Horizon 1):

Variates X11 X2 X3 X4 X5 X6 X9 X10
Means 3:31 3-39 1:22 0-96 2-84 0-96 1-33 1-08
Sample size29 30 30 29 IM I A DW
Variance-
covariance 0:50 0-39 0-21 0-10
Matrix 0:57 0-14 0-08
0-18 0-05
0-05

0-23
0-36
0-09
0-06
0-39

0-04
0-04
0-03
0-01
0-03
0-02

0-20
0-14
0-13
0-04
0-08
0-03
0-14

0-14
0-14
0-08
0-03
0-08
0-02
0-08
0-07

Brachial valve internal moulds (Kildare, Horizon 1):

Variates XX XGA SK 4

Means 3:07 3-44 1-13 0-87 0-77 1-28 2-54
Sample size i) Su. «SS 3S) SB) BE

Variance-

covariance 0:46 0:38 0:14 0:08 0-08 0-10 0-33

Matrix 0:76 0:09 0:10 0:08 0-15 0:26

0-10 0-01 0-03 0-02 0-01

0-04 0-01 0-03 0-06

0-03 0:03 0-06

0-06 0-08

0.30

DISCUSSION. The numerical data in the description are based
only on the mean values for the large sample from Grange
Hill, Kildare, where this species is a very common element of
the fauna. The poor preservation of the samples from Kil-

igs 1-3 Platystrophia sp. 1. Kildare, Grange Hill Horizon 1. 1, BC 12677, detail of ornament of latex cast, x 10. See also Pl. 4, figs 12, 15. 2,
BC 12675, detail of ornament of latex cast showing the accentuated lamellae, x 10. See also PI. 4, fig. 10. 3, BC 12681, internal mould of

igs 4-5 Platystrophia sp. 2. Carrigadaggan. BC 12682, internal mould of brachial valve and latex cast, x 4.
‘igs 6-11 Rhactorthis sp. Kildare, Grange Hill Horizon 1. 6, BC 12683a, internal mould of brachial valve, x 4. 7, BC 12684, external mould

16, x 4

x 10.

| of pedicle valve, x 4. 8, 9, BC 12685a, ventral and dorsal views of a conjoined internal mould, x 4. 10, BC 12685b, latex cast of external
mould of conjoined valves, counterpart of Figs 8-9, dorsal view, x 4. 11, BC 12683b, external mould of brachial valve, counterpart of Fig.

igs 12-16 Cremnorthis parva Williams. Kildare, Grange Hill Horizon 1. 12, BC 12686, latex cast of external mould of brachial valve, x 10.
13, 14, BC 12687, external mould and latex cast of brachial valve, x 10. 15, 16, BC 12688, external mould and latex cast of brachial valve,

146

LAURENT UA

OCEAN

1APETUS

Avalonia (after Parkes, 1992).

bride and Carrigadaggan made measurement of all variates
difficult, but a principal component analysis (PCA) of all
three samples shows that in plots of the first four eigen-
vectors (see Fig. 17) the two small samples fall within the
same region as the Kildare sample. It is possible that analysis
of a larger topotype sample of the species would show
significant differences between the Welsh and Irish forms,
but the original description was based on fewer than ten

ANGLO-WELSH

GONDWANALAND i

Fig. 16 Palaeogeographic reconstruction of the Iapetus region in mid-Caradoc times (c. 448 Ma) showing the main continental masses of
Laurentia, Baltica, Gondwana and Eastern and Western Avalonia. The main brachiopod provinces of Scoto-Appalachian, Baltic and
Mediterranean affinity are also shown. At this time, an Anglo-Welsh Province, including these Leinster faunas, was centred on Eastern

SCOTO-APPALACHIAN
PROVINCE

_ ‘BALTIC
PROVINCE

EAST AVALONIA

valves. The mean percentage length of the ventral muscle
field relative to the valve length is significantly longer in the
Irish form (Kildare — 39% compared to 35%, p < 0-01). This |
is considered inadequate to justify erection of even a new
subspecies, since the Bala sample consisted of only 5 speci-
mens. A larger, better preserved sample would probably

encompass the same variation seen in the Kildare form. f

PLATE 6

Figs 1-7 Cremnorthis parva Williams. Figs 1-5, Kildare, Grange Hill Horizon 1. 1, BC 12689, internal mould of brachial valve, x 5. 2, 3, BC
12690, internal mould of pedicle valve, latex cast and mould, x 15. 4, BC 12691, internal mould of brachial valve, and BC 12692, internal
mould of pedicle valve, both x 6. 5, BC 12693, internal mould of brachial valve, x 3. Figs 6, 7, Carrigadaggan. 6, BC 12694, ventral view
of conjoined internal mould, x 10. 7, BC 12695, internal mould of pedicle valve, x 10.

Figs 8-16 Skenidioides costatus Cooper. Kildare, Grange Hill Horizon 1. 8, 12, BC 12696, dorsal and ventral views of conjoined internal
mould, x 10. 9, BC 12697, internal mould of brachial valve, x 10. 10, BC 12698, internal mould of brachial valve, x 10. 11, BC 12699,
ventral view of conjoined internal mould, x 10. 13, BC 12700, internal mould of pedicle valve, x 10. 14, BC 12701, internal mould of
pedicle valve, x 10. 15, BC 12702, internal mould of pedicle valve,

x 10. 16, BC 12703, internal mould of pedicle valve, x 10.

3RACHIOPODS OF DUNCANNON GROUP OF SE IRELAND 147

148

Family SKENIDIIDAE Kozlowski, 1929
Genus SKENIDIOIDES Schuchert & Cooper, 1931

Skenidioides costatus Cooper, 1956
Pl. 6, figs 8-16; Pl. 7, figs 1-5
1956 Skenidioides costatus Cooper: 493; pl. 97, figs
38-48.
aff. 1962 Skenidioides aff. costatus Cooper; Williams: 126;
pl. 11, figs 24-27, 52.

cf. 1963 Skenidioides cf. costatus Cooper; Williams:
375-377; pl. 4, figs 7-14.
cf. 1974 Skenidioides cf. costatus Cooper; Williams:

82-83; pl. 13, figs 14-16; pl. 14, figs 1-3.
cf. 1979a Skenidioides cf. costatus Cooper; Hurst: 242; figs
145-159.

MATERIAL AND LOCALITIES. Kilbride: 2 internal and 3 exter-
nal moulds of pedicle valves; 3 internal and 6 external moulds
of brachial valves. Kildare, Grange Hill, Horizon 1: 10
internal and 10 external moulds of brachial valves; 19 internal
and 3 external moulds of pedicle valves. Kildare, Grange Hill
House Cottage: 2 internal and 1 external moulds of pedicle
valves; 1 internal and 2 external moulds of brachial valves.
Carrigadaggan: 1 internal mould of a pedicle valve. Kildare,
Grange Hill, Horizon 2: 3 internal moulds of pedicle valves.
Greenville-Moyne: 1 internal and 1 external moulds of a
pedicle valve.

DESCRIPTION. Exterior. Ventribiconvex, subpyramidal Sken-
idioides with pedicle valve length about seven-tenths of the
length, and about 40% as deep as long. Brachial valve gently
convex with distinct median sulcus, about 50-70% as long as
wide. Ornament of radial costellae, about 2-5, commonly 3,
ribs per mm 2mm anteromedially of umbo, in both valves.
Commonly a wider median rib on the pedicle valve, with total
rib counts of between 12 and 20 with 15-17 the most common
frequency. Ventral interarea high, catacline to apsacline with
open delthyrium. Dorsal interarea shorter, anacline.

Ventral interior. Generally unsupported spondylium about
a quarter as long as the valve, and about 94% as long as wide.
Some shells have a median thickened ridge of shell support-
ing the spondylium.

Dorsal interior. Thin median septum, continuous anteriorly
from shaft-like cardinal process, extending about 90% of
valve length. Slender brachiophores with bases convergent
onto median septum defining a diamond shaped cruralium
about a third as long as valve.

DISCUSSION. Previously described samples compared to S.
costatus (Cooper 1956) differ in some proportions from each
other and from the material described here, but the differ-
ences are not considered important enough to warrant taxo-
nomic recognition. Principal component analysis of all the

PLATE 7

M. A. PARKES ||

MEASUREMENTS
Pedicle valve internal moulds (Kildare, Horizon 1):

Variates Xl X2 x4 XG eel 4
Means 3:37 4-64 1:29 0-81 0-84
Sample size 19 19 I) 18 18
Variance-covariance 0-46 0-42 0:06 0-13 0-06
Matrix 0:77 0-05 0-10 0-08
0-08 0-01 0-01
0-06 0-03
0-02 °
Brachial valve internal moulds:
Variates X1 X2 X15 X16 X24
Means 3:00 4-11 0-98 0-83 2-76
Sample size 10 10 10 10 10
Variance-covariance 0:25 0-24 0:06 0-02 0-26
Matrix 0:78 0-02 0-10 0-14
0-03 0.0 0.06
0-02 0-0
0-32
Brachial valve external moulds:
Variates Xl X2 x4
Means 2:73 5-32 10-47
Sample size 10 10 7
Varlance-covariance 0-43 0-63 ~=0-05
Matrix 1A S014)
0-12

material shows no differentiation on the first four vectors, as
shown in Fig. 18. As noted by Mitchell (1977: 50) and Hiller
(1980: 146), internal features of many Skenidioides species
show no significant differences, the species being separated
on the basis of the density of external ornament. The rib
counts for this Irish material is comparable to previously
described populations of S. costatus of similar size range.
One aspect which apparently requires further investigation
is the branching mode of the ribs of the species. Williams
(1974: 83) discussed the differences between Shelve and Bala
stocks, with new costellae arising only from the ventral
median rib in the former and branching freely from lateral
costae in the latter. In the Bala stocks, the majority of costae
branched externally (Williams 1963: 377). However, Hurst
(1979a: 242) states that the Shropshire stock only branched

Figs 1-5 Skenidioides costatus Cooper. Kildare, Grange Hill Horizon 1. 1, 3, BC 12704, external mould of brachial valve and interareas, x 6,
and enlarged oblique posterior view of latex cast of interareas, showing hinge line and open delthyrium and notothyrium, Xx 10. 2, BC
12809, latex cast of external mould of pedicle valve, x 6. 4, 5, BC 12705, external mould of brachial valve and interareas, and latex cast

showing open delthyrium, x 10.

Figs 6-11 Oanduporella cf. reticulata Hints. Kildare, Grange Hill House Cottage. 6, 9, BC 12706, external mould of pedicle valve, latex cast
and mould, x 10. 7, 8, BC 12707a (upper) and BC 12708a (lower); internal moulds and latex cast of 2 pedicle valves, x 4. 10, BC 12711b,
external mould of pedicle valve, x 4. 11, BC 12711a, internal mould of pedicle valve, counterpart of Fig. 10, x 4.

Fig. 12 View of slab, containing BC 12670, external mould of Plaesiomys multiplicata Bancroft (P; see also Pl. 4, figs 3-4), to illustrate the
typical assemblage at Kildare, Grange Hill House Cottage; with Oanduporella cf. reticulata Hints (A), Rafinesquina sp. (B), and

numerically dominant gastropods (C). x 1.

;

p
4sRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND 149

150
V-2
-5.1 -3.3 -1.5 0.3 2.1 3.9
V-1 C24V2
0.9
0.5
0.1
V-4

-0.3

-0.7

-0.5 0.3 1.1 1.9
V-3 C24V2

$2511” Fibs)

Fig. 17 Principal component analysis of samples of Cremnorthis
parva, internal moulds of pedicle valves only. Top, vector 1
against vector 2. Below, vector 3 against vector 4. VW =
Carrigadaggan, * = Kilbride, 0 = Kildare, Grange Hill Horizon
il.

internally. The present material has few external moulds with
costellae, but those that do, show both internal and external
branching.

M. A. PARKES |

Superfamily ENTELETOIDEA Waagen, 1884
Family DALMANELLIDAE Schuchert, 1913
Genus OANDUPORELLA Hints, 1975 |

Oanduporella cf. reticulata Hints, 1975
Pl. 7, figs 6-12; Pl. 8, figs 1-7 |

cf. 1975 Oanduporella reticulata Hints: 19, 105; pl. 1, figs
1-15; pl. 2, figs 1-5. |

1980a? Ravozetinal/Onnizetina; Mitchell, in Romano: 206.
1985 Oanduporella cf. reticulata Hints; Harper & Mitch-
ell, in Harper et al.: 295, figs 25-37.

MATERIAL AND LOCALITY. Kildare, Grange Hill House Cot- |
tage: 10 internal and 6 external moulds of brachial valves, 13 |
internal and 9 external moulds of pedicle valves.

DISCUSSION. Harper & Mitchell gave a full description |
(Harper et al., 1985) of material they compared to Hints’
species from the east Baltic, with which the present material -
from Kildare accords well. The Herbertstown material, from '
the Clashford House Formation, was the first record of the |
genus from Britain or Ireland and the present sample repre- |
sents the second known occurrence from these areas. It
serves to emphasize the similarities of the Kildare fauna to |
that from Herbertstown with two conspecific forms, Plaesi-
omys multiplicata Bancroft and Oanduporella cf. reticulata |
Hints, present, with a possible third, Hibernodonta? Harper |
& Mitchell (in Harper et al. 1985). There are few suitable —
specimens but similar rib counts (5-6 per 2 mm at 5 mm_
sagittally) are seen in the Kildare specimens, although the |
microsculpture is well developed on most specimens.

Oanduporella sp. (Not figured) .

MATERIAL AND LOCALITY. Greenville: 3 external and 1
internal moulds of pedicle valves, 1 external mould of a)
brachial valve. .
DISCUSSION. The poor preservation, and deformation in this
mudstone lithology made it impossible to compare this mate-
rial to the specimens from Kildare, Grange Hill House |
Cottage. Nevertheless, this small sample shows the character-_
istic pitted microsculpture of the genus and extends the
known geographical range. |

Genus REUSCHELLA Bancroft, 1928

Reuschella ? sp. Pl. 11, fig. 16

|
|
|
|

MATERIAL AND LOCALITY. Kilbride: A single internal mould |
of a pedicle valve.

DISCUSSION. The single poorly preserved mould is assigned |
to Reuschella on the basis of the sharp median carina, curved
long apsacline interarea, the ventral muscle scar and massive
teeth. The specimen represents the sole occurrence of the
genus within the southeast Ireland Caradoc. It is known from
the Actonian of Shropshire (Hurst 1979a), the Soudleyan of
Shelve, Shropshire (Williams 1974) and Bala (Williams 1963)
as well as from Girvan (Williams 1962).

e
;

3RACHIOPODS OF DUNCANNON GROUP OF SE IRELAND 151

0.9

0.5
|
0.1
. V4
|
-0.3
-0.7
-1.1
-214 -1.1 ft) 09 19 2.9
V-3 SKENPV
B
2.9
1.9
0.9
v2

—2.1 One W073 1.5 Pet eye) -21 -0.9 0.3 1.5 27 ~3t9
V-1 SKBVEX V-1 SKBVEX

Ea 199 087 0.5 Ai” 72:9 -3.1 -19 -0.7 0.5 i729
| V-1 SKEEXT V-1 SKEEXT

lig. 18 Principal component analysis of Skenidioides costatus, from Kilbride (@) and Kildare, Grange Hill House Cottage (*) and Horizon 1
, (Q). A, pedicle valve internal moulds; B, brachial valve external moulds; C, pedicle valve external moulds.

152

Family LINOPORELLIDAE Schuchert & Cooper, 1931
Genus SALOPIA Williams, in Whittington & Williams 1955
Pl. 11, figs 18-20

MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon
2: a single internal mould of a pedicle valve. Greenville-
Moyne: 4 internal and 1 external moulds of brachial valves.

Salopia sp.

DESCRIPTION. Ventral valve. Interior, strongly convex
mould, slightly sulcate, over half as deep as long, and slightly
wider than long. Maximum width just anterior to hinge line,
giving a subcircular outline to shell with high apsacline, but
nearly catacline interarea. Short teeth are supported by
divergent dental plates. Muscle field extends beyond dental-
plates to about one-third of valve length. External ornament
only seen where it is impressed around margins of shell
interior.

Dorsal valve. Interior, gently convex in mould form, with
low notothyrial platform between divergent brachiophores
carrying a thin linear shaft which is continuous with a low
median septum, extending to about mid-length. External
ornament impressed slightly around margins of interior.

MEASUREMENTS (mm). BC 12758: X1 = 11-5, X2 = 12, X4
= 6, X9 = 4-5 (Pl. 11, fig. 19).

DiscussION. Salopia is known from the Llandeilo (Lockley
& Williams 1981: 51) and Lower Caradoc (Williams 1963,
1974; Whittington & Williams 1955) of Wales and Shrop-
shire. The single pedicle valve from Kildare is inadequate for
formal comparison, but is apparently significantly deeper
than described species. The sample from Greenville-Moyne is
indistinctly preserved, and is lacking in pedicle valves, so
cannot be directly compared to the Kildare specimen. The
closest obvious comparison is with Salopia salteri (Davidson,
1869).

Family SAUKRODICTYIDAE Wright, 1964
Genus SAUKRODICTYA Wright, 1964

Saukrodictya cf. sp. A of Hints Pl. 8, figs 8-15

cf. 1979 Saukrodictya sp. A, Hints: 57; pl. 2, fig. 10; pl. 4,
figs 15-22.

MATERIAL AND LOCALITIES. Carrigadaggan: 1 internal and 3
external moulds of pedicle valves, 4 external moulds of
indeterminate valves. Greenville-Moyne: 2 indeterminate
external moulds.

DESCRIPTION. Exterior. Typical ornament of exopuncta, up
to 6 radial rows, closely spaced, in the interspaces between
narrow ribs. Interspaces are relatively wide and rounded.
Ventral valve gently convex, wider than long.

Ventral interior. Nearly 50% wider than long. Gently
convex profile, rectimarginate commissure. (Dorsal valve
unknown).

M. A. PARKES jh sh

DIscussION. The mould material, although very poor, 1s |
assigned to Saukrodictya rather than Salacorthis because of |
the typical pitted ornament found only in the interspaces and
not on the thin ribs. The frequency of ribs is also greater than
in Salacorthis costellata Williams (1974), the only known
species, and the pedicle valves described here are not sulcate,
as are those of Salacorthis.

There are a number of described species of Saukrodictya,
but in all cases they are based on limited material and are not
well known. The present material does not permit a detailed
comparison with described species, but the illustrations of
Saukrodictya sp. A by Hints (1979) from the Idavere and
Johvi Stages in Estonia (L. Caradoc — multidens Biozone) are
most similar to this material. The species apparently lacks the
strong fold of S. reticula Vinassa, 1927 (Villas, 1985). It has a —
lower frequency of ribs than S. rotundopora Hints or S. |
oblongatopora Hints, both of which also have a fold. Simi- |
larly, S. porosa is sulcate and also has a greater rib frequency,
though its general outline is similar (Havlicek 1977). The type |
species S. hibernica Wright (Wright 1964; see also Hiller, |
1980) is strongly sulcate. However, Villas (1985) has sug-
gested that S. hibernica may be conspecific with S. reticula
(Vinassa, 1927), ‘but there are too many gaps in the knowl-
edge of ‘British‘ and Sardinian Saukrodictyae’. The present —
sample unfortunately does nothing to clarify the definition of
species, but is stratigraphically and biogeographically signifi- —
cant.

These are the oldest known occurrences of Saukrodictya in
Ireland. S. rotundopora Hints (1979: 53) and S. oblongato-
pora Hints (1979: 55) are from approximately contemporane-
ous stages in the Middle Caradoc of Estonia. Other
occurrences are Ashgill in age, including the type species S.
hibernica from Portrane (Wright 1964: 216) and Wales (Hiller
1980: 165), S. wrighti from Belgium (Sheehan 1987) or from |
the Llandovery (S. sp. from Wales (in Temple, 1970: 32); S. .

|
H
|

sp. B from Estonia (Hints, 1979: 58)). According to Havlicek
(1977) the oldest occurrence of species of Saukrodictya are S.
porosa from the Liben and Letna Formations of Bohemia |
(Middle Llandeilo to Costonian) and in the Costonian/ |
Harnagian of Portugal (Mitchell 1974). It would thus appear
to have a Gondwanan origin and to have migrated north: |
ward, reaching Ireland by the Longvillian or earlier.

Superfamily GONAMBONITOIDEA Schuchert & Cooper, )
1931

Family KULLERVOIDEA Opik, 1934 |

Genus KULLERVO Opik, 1934

Kullervo aff. hibernica Harper, 1952
1218

aff. 1952 Kullervo hibernica Harper: 100; pl. 6, figs 6-8.
aff. 1977 Kullervo aff. hibernica Harper; Brenchley et al.: |
70. |

PI. 9, figs 1-10,

PLATE 8

Figs 1-7 Oanduporella cf. reticulata Hints. Kildare, Grange Hill House Cottage. 1, 2, BC 12709, external mould of brachial valve, and latex
cast, X 10. 3, 4, BC 12710a, internal mould of brachial valve, and latex cast, x 10. 5, 6, BC 12710b, external mould of brachial valve,
counterpart of Figs 3—4, latex cast, x 10. 7, BC 12712, internal mould of brachial valve, x 72.

Figs 8-15 Saukrodictya cf. sp. A of Hints. Carrigadaggan. 8, 11, BC 12713, external mould, latex cast and mould, x 10. 9, 10, BC 12714,
external mould and latex cast, x 10. 12, BC 12715a, internal mould of pedicle valve, x 8. 13, BC 12716b, latex cast of external mould, x
10. 14, 15, BC 12715b, external mould of pedicle valve, counterpart of Fig. 12, latex cast and mould, x 10.

}

oa)
w

3RACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

M. A. PARKES

154

i
)

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

MATERIAL AND LOCALITIES. Carrigadaggan: 3 internal
moulds and 5 external moulds of pedicle valves, 2 internal
and 3 external moulds of brachial valves and 8 indeterminate
‘external moulds. Kildare, Grange Hill, Horizon 1: 1 external
mould of a pedicle valve and2 fragments of internal moulds of
pedicle valves. Greenville-Moyne: 3 internal and 3 external
moulds of pedicle valves, 2 external moulds of brachial
valves, and 2 indeterminate external moulds.

DESCRIPTION. Exterior. Strongly ventribiconvex Kullervo
with pyramidal pedicle valve and maximum width along hinge
line. Concave posterolateral flanks on extended hinge line,
anterior slope evenly convex. Shallow median sulcus in
brachial valve, but essentially rectimarginate commissure.
Ventral interarea high, curved, apsacline near umbo, cata-
cline near hinge. Dorsal interarea short, anacline. Ornament
of distinctive reticulate pattern, of strongly developed con-
centric lamellae and regular radial ribs. Radial ornament
absent on posterolateral flanks.

| Ventral interior. Spondylium with hemisyrinx supported by
well-developed median septum extending nearly to mid
length. External ribs impressed on anterior margins of valve
interior.

| Dorsal interior. Cardinalia of thick divergent socket ridges
about three times as long as wide, extending less than half
valve width. Thin cardinal process in narrow space between
their ends. Socket ridges merge anteromedially with thick
notothyrial platform, itself passing into thick median ridge
anteriorly. At about mid-length of valve the ridge tapers to a
thin, low median septum, separating very poorly impressed
adductor scars.

Discussion. Harper’s (1952) original description of Kullervo
hibernica was based on limited material. In respect of the
external ornament, overall shape and ventral interior the
described material is comparable to the paratype material of
Kullervo hibernica Harper from Grangegeeth (NMING:
F14035, 14036). However, a single well-preserved brachial
valve interior from Carrigadaggan (PI. 9, figs 12, 13) shows
some differences from the holotype of hibernica (NMING:
\F14034; Pl. 9, fig. 18), which is itself broken and missing
posteriorly; in the hibernica holotype the impression of
external ribs is more regular, stronger and abruptly and
evenly terminated. In the Carrigadaggan specimen the ribs
fare variably impressed, also more irregular and longer.
However, an additional broken, poor specimen does show a
more reg<iar and even impression of external ribs. In both
specimens, a thick, well-developed median ridge, continuous
with the notothyrial platform, tapers at about mid-length to a
thin, low median septum between the anterior adductors.
The most noticeable difference is in the anterior adductor
sears. In K. hibernica they are very strongly impressed with

PLATE 9

155

raised edges, but in the Carrigadaggan specimen they are
barely seen and less divergent.

With such a small sample these differences are not deemed
to justify erection of a new species. Larger samples are
needed to clarify the relationship of the Irish specimens of
Kullervo to each other and other poorly known species,
especially since the material described herein is mostly frag-
mentary or broken. It is thus difficult to measure the impor-
tant morphological characters, but the figured specimens
show similar proportions to the type material. Other recent
descriptions of Kullervo species, such as Whittington &
Williams (1955) from the Derfel Limestone of Wales, Wright
(1964) from the Portrane Limestone of eastern Ireland, and
Hiller (1980) from North Wales, have all drawn attention to
Opik’s original description of the genus (1934). In this, he
defined four groups on the basis of external ornament. All
the present material is similar to group 2, in particular K.
lacunata Opik, which has strong radial ribs in the middle
sector but dominant concentric ornament on the ears. The
material is unlike K. complectens albida which has a ventral
sulcus bounded by stronger ribs. Consequently, although the
type material is poorly known, this material is affiliated to K.
hibernica. It is possible that larger collections of topotype
material may show that K. hibernica is synonymous with the
Kukruse (N. gracilis) form from Estonia, K. lacunata, or
more likely that it is descended from that species, with an
increase of radial ribs and stronger reticulation.

Superfamily TRIPLESIOIDEA Schuchert, 1913
Family TRIPLESIIDAE Schuchert, 1913
Genus BICUSPINA Havli¢ek, 1950

PI. 9, fig. 11

MATERIAL AND LOCALITY. Kilbride: 2 internal moulds of
brachial valves, and one external mould fragment.

Bicuspina ?sp

DISCUSSION. These two incomplete dorsal moulds could pos-
sibly be assigned to Bicuspina, or to the similar genus
Oxoplecia which Carlisle (1979: 552) recorded from Kilbride.
However, the overall shape, especially the very angular
dorsal fold, suggests Bicuspina is more appropriate.

Superfamily PLECTAMBONITOIDEA Jones, 1928
Family BIMURIDAE Cooper, 1956
Genus BIMURIA Ulrich & Cooper, 1942

Bimuria cf. dyfiensis Lockley, 1980
Pl. 9, figs 14-17; Pl. 12, figs 1-9
21977 Bimuria sp.; Mitchell: 95; pl. 19, figs 24-28.

Figs 1-10, 12-13 Kullervo aff. hibernica Harper. Carrigadaggan. 1, BC 12717, incomplete internal and external mould of pedicle valve, x 4.
2, BC 12718a, internal mould of pedicle valve, x 2. 3, 7, BC 12719, external mould of pedicle valve, and latex cast, x 2. 4, 5, BC 12720,
internal mould of pedicle valve, and latex cast, x 4. 6, BC 12721, latex cast of external mould of pedicle(?) valve, x 4. 8, BC 12718b, latex

| cast of external mould of pedicle valve, counterpart of Fig. 2, x 4. 9, 10, BC 12722, external mould of brachial valve, latex cast and mould,
x 4. 12, 13, BC 12723, internal mould of brachial valve, latex cast and mould, x 4.

Fig. 11 Bicuspina? sp. Kilbride. BC 12724, internal mould of brachial valve, x 2.

[Figs 14-17 Bimuria cf. dyfiensis Lockley. Ballykale. 14, BC 12725, internal mould of pedicle valve, x 32. 15, BC 12726, dorsal view of
internal mould of pedicle valve showing strongly incurved umbo, x 32. 16, BC 12727a, dorsal view of conjoined internal mould, x 32. 17,

BC 12728, internal mould of pedicle valve, x 32.

\Fig. 18 NMING:F14034. Holotype of Kullervo hibernica Harper; an internal mould of a brachial valve from Grangegeeth, Co. Meath. x 4.

156
cf. 1980 Bimuria dyfiensis Lockley: 215, figs 60-62, 64-65.

MATERIAL AND LOCALITY. Ballykale: 16 internal moulds of
pedicle valves, 3 internal moulds of brachial valves and 4
external moulds of pedicle valves.

DESCRIPTION. Exterior. Concavo-convex valves with pedicle
valve umbo incurved and overlapping dorsal interarea. Recti-
marginate, with subcircular outline. Both valves essentially
smooth, with comae absent or very indistinct, resembling
growth lines, in the brachial valve. Mean length about
three-quarters of width.

Ventral interior. Simple teeth developed laterally for about
one-third of the width of the valve but short anteromedially.
A pinnate mantle canal pattern surrounding an undifferenti-
ated muscle field is variably impressed (or preserved).

Dorsal interior. Low socket ridges nearly parallel to the
hinge line. Prominent submedian septa and a thin median
septum within a papillose bema.

MEASUREMENTS
Pedicle valve internal moulds:

Variates X1 X2
Means 9-89 13-3
Sample size 16 16
Variance-covariance 12-21 11-36
Matrix 15-55

DISCUSSION. Specimens are too deformed for reliable quan-
titative study. The measurements taken are given above, but
caution is advised in using them other than as a general guide
to the species morphology, because of tectonic deformation.
The Ballykale population of Bimuria is quite strongly
deformed and flattened, so precise comparison with known
Scoto-Irish species (B. cf. buttsi Cooper, B. youngiana
Davidson, B. youngiana recta Williams) is not possible.
Enough well-preserved specimens are described to justify
assignment to B. cf. dyfiensis; the most significant feature of
the species is the absence of comae. The size of the Ballykale
sample is similar to the Welsh sample described by Lockley
(1980) from the Gelli-grin Formation of the Bala area of
Wales. The lack of a fold and sulcus is distinctive in this
population. Bimuria sp. from Kilbride (below) is of a much
smaller mean size but otherwise shows very little difference
from B. cf. dyfiensis. It is described under open nomenclature
in the absence of dorsal interiors and exteriors, and because it
is also similar in internal morphology and size to B. youngi-
ana recta Williams, One poor pedicle valve exterior, how-
ever, shows no evidence of comae. The lack of comae is a
feature of Bimuria sp. from the Ashgill Killey Bridge Forma-
tion at Pomeroy (Mitchell 1977), although that form is gently

PLATE 10

Figs 1-7 Bimuria sp. Kilbride. 1, BC 12729, internal mould of pedicle valve, x 4. 2, BC 12730, internal mould of pedicle valve, x 4. 3, BC
12731, internal mould of pedicle valve, x 4. 4, BC 12732, internal mould of pedicle valve, x 4. 5, BC 12733, internal mould of pedicle
valve, x 4. 6, BC 12734a, internal mould of pedicle valve, x 4. 7, BC 12734b, external mould of pedicle valve, counterpart of Fig. 6, x 4.

Figs 8-14 Leptellina (Leptellina) cf. llandeiloensis (Davidson). Carrigadaggan. 8, 9, BC 12735, internal mould of pedicle valve, and latex cast,
x 4. 10, BC 12736, internal mould of pedicle valve, x 2. 11, 13, 14, BC 12737, internal mould of pedicle valve, posterior view, latex cast,
and dorsal view, x 2. 12, BC 12738, latex cast of internal mould of brachial valve, x 4.

Figs 15-21 Leptestiina oepiki Whittington. Figs 15-18, Kildare, Grange Hill Horizon 1. 15, BC 12739, internal mould of pedicle valve, x 4.
16, 17, 18, BC 12740, internal mould of brachial valve, posterior and dorsal views of latex cast, and mould, x 4. Figs 19-21, Kilbride. 19,
BC 12741, internal mould of brachial valve, x 4. 20, 21, BC 12742, internal mould of brachial valve, latex cast and mould, x 4.

|
i
M. A. PARKES}
sulcate. The size of specimens in this small sample is interme-|
diate between B. cf. dyfiensis and Bimuria sp. from Kilbride.
It seems likely that all three samples are closely related, but!
further specimens, preferably undeformed, are needed to’
verify this.

Bimuria sp. PI. 10, figs 1-7
|

MATERIAL AND LOCALITY. Kilbride: 21 internal moulds and
one external mould of pedicle valves.

DESCRIPTION. Exterior. Unknown, except for one smooth,
but poorly preserved valve. 2

Ventral interior. Strongly concavo-convex valve with strongly |
incurved umbo of pedicle valve overlapping dorsal interarea.
Rectimarginate anterior commissure. Shell outline variable,
from occasionally subcircular to sometimes transverse. Maxi- |
mum width just anterior of hinge line, at about 27% of length.
Length is always less than width; mean value is 78%. Depth of
ventral valve is nearly half the length (44%).

Dorsal interior. Unknown.

MEASUREMENTS
Pedicle valve internal moulds (Kilbride):

Variates X1 x2 x3 xX4 |
Means 6:12, 7-90 Sales 2-71 |
Sample size 19 20 20 20
Variance-covariance 2:62 2:51 0-89 1:16 |
Matrix 3:23 0-65 1:01 |
0-69 0-44
0-81

DISCUSSION. See Bimuria cf. dyfiensis, above. |

Family LEPTELLINIDAE Ulrich & Cooper, 1936
Subfamily LEPTELLININAE Ulrich & Cooper, 1936
Genus and subgenus LEPTELLINA (LEPTELLINA) Ulrich
& Cooper, 1936

Leptellina (Leptellina) cf. llandeiloensis (Davidson, |
1883) Pl. 10, figs 8-1

cf. 1883 Leptaena llandeiloensis Davidson: 171; pl. 12, fig.
26, non figs 27-29.

cf. 1917 Plectambonites Llandeiloensis (Davidson); Reed:
876; pl. 13, figs 32-34; pl. 14, figs 1-3.

cf. 1928 Leptelloidea llandeiloensis (Davidson); Jones: 477.

cf. 1962 Leptellina llandeiloensis (Davidson); Williams: 164;)
pl. 15, figs 27-29, 32.

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

158

cf. 1977 Leptellina cf. llandeiloensis (Davidson); Mitchell:
72; pl. 13, figs 14-17.
cf. 1978 Leptellina llandeiloensis (Davidson); Cocks: 93.

MATERIAL AND LOCALITY. Carrigadaggan. 3 internal moulds
of pedicle valves, and 3 incomplete internal moulds of
brachial valves.

DESCRIPTION. Exterior. Unknown.

Ventral interior. Convex, transverse, nearly semicircular
about 70% as long as wide and 40% as deep as long,
rectimarginate anterior commissure. Maximum depth at
about mid-length. Orthocline interarea about 20% of valve
length. Deeply impressed large, quadrilobate muscle field
80% as long as wide, and extending anteriorly to about
mid-length of valve. Small diamond-shaped platform anterior
of and between the muscle field lobes, depressed in centre
and with 4, 6 and 8 coarse pustules in a row on the anterior
slope of the platform, sagittally. Delthyrium apparently open.
Saccate pattern of mantle canals.

Dorsal interior. Large, well-defined platform is strongly
elevated, ankylosed with a median septum and medially
indented.

DiscussION. The present material is clearly inadequate for
an unequivocal identification, in the absence of complete
brachial valve interiors or any exteriors, but is assigned to L.
cf. llandeiloensis because of the strong similarities to the
material described by Mitchell (1977: 72) from the Caradoc
Bardahessiagh Formation of Pomeroy. Previous descriptions
have not recorded or remarked on the presence of a discrete
median row of pustules anteriorly of the diamond shaped
platform, but Mitchell’s (1977: pl. 13, figs 14, 16) figured
specimens clearly show them. What significance should be
attached to this is unknown, since although Williams’ (1962:
pl. 15, fig. 28) specimens show them, he also figures a
specimen of Leptellina semilunata (1962: pl. 15, fig. 23) which
has a less ordered but equally strong row.

Family LEPTESTIIDAE Opik, 1933, emend. Cocks & Rong,
1989
Genus LEPTESTIINA Havlicek, 1952

REMARKS. Cocks & Rong (1989: 116) reduced Leptestiina to
a subgenus of Leangella Opik, 1933. This is not followed here
since, although Melou (1971) showed a_ phylogenetic
sequence from Leptestiina derfelensis through L. prantli and
L. aonensis to Tufoleptina (=Leangella), the early members
of the lineage such as Leptestiina oepiki, L. derfelensis and L.
indentata are sufficiently distinctive from Leangella, in the
lack of a platform (sensu Cocks & Rong, 1989) anteriorly.

M. A. PARKES| |!

Leptestiina oepiki Whittington, 1938
Pl. 10, figs 15-21; Pl. 11, figs 1-6

1938 Sampo oepiki Whittington:255; pl. 10, figs 15-16; pl,
11, fig. 10.

1963 Leptestiina oepiki (Whittington); Williams: 428-430; |
pl.10, figs 15, 16, 19-21.

1978 Leptestiina oepiki (Whittington); Cocks: 94.

1989 Leangella (Leptestiina) oepiki (Whittington); Cocks &|
Rong: 116-117.

MATERIAL AND LOCALITIES. Kilbride: 6 internal and 1 exter-|
nal moulds of brachial valves; 4 internal and 1 external mould’
of pedicle valves. Kildare, Grange Hill, Horizon 1: 4 internal
moulds of a pedicle valve and 2 internal moulds of brachial]
valves. Greenville-Moyne: 3 internal moulds of pedicle valves’
and 2 internal moulds of brachial valves.

DESCRIPTION. Exterior. Evenly concavo-convex valves, of
semicircular outline. Pedicle valve about 60% as long as’
wide, and about 35% as deep as long. Ornamentation poorly-
known, unequally parvicostellate with very fine costellae
separated by few thicker ribs. Ventral interarea apsacline,
dorsal interarea hypercline. |

Ventral interior. Short blunt teeth supported by short!
receding dental plates, bordering a transversely subpentago- |
nal muscle field which extends about 23% of valve length —
anteriorly. Length of muscle field about 50% of width.
Deeply impressed vascula markings with lemniscate pallial
sinus pattern.

Dorsal interior. Characteristically longitudinally and radi-.
ally striated bema, bilobed and undercut with median incision
and septum separating the two rounded lobes which originate
laterally from the socket ridges, bounding oval sockets. Bema
is about two-thirds as long as wide and extends anteriorly half’
the length of the valve.

Discussion. The species L. oepiki is known from the |
Longvillian of Wales (Williams 1963: 428) and possibly the
Actonian of Shropshire (Hurst 1979a: 275). Williams (1963:
430) noted the similarities between L. oepiki and the closely |
related Costonian species L. derfelensis Jones, 1928, from the
Derfel Limestone in North Wales. He suggested the two may
be synonymous, but the resolution of this must await further
collections of Leptestiina derfelensis as well as move Irish
material. The present sample is too small and poorly pre- |
served to assess the variability of the species, particularly the
external ornament. L. aff. oepiki is also known from
Pomeroy, Co. Tyrone, in the Cautleyan Killey Bridge Forma-
tion (Mitchell 1977: 76); this also resembles L. derfelensis in
some respects.

Harper (in Harper & Owen 1984: 29) revised the Upper |
Caradoc Norwegian species L. indentata (Spjeldnaes 1957: |

PLATE 11

Figs 1-6 Leptestiina oepiki Whittington. Kilbride. 1, BC 12743, internal mould of pedicle valve, x 4. 2, 3, BC 12744, internal mould of
brachial valve, and latex cast, x 4. 4, BC 12745, internal mould of pedicle valve, x 4. 5, BC 12746, internal mould of brachial valve, x 4.

6, BC 12747, internal mould of brachial valve, x 4.

Figs 7-15, 17 Leptestiina oepiki ampla subsp. nov. Carrigadaggan. 7, 8, Holotype BC 12748, internal mould of brachial valve, and latex cast, Xx
4. 9, 10, BC 12749, internal mould of brachial valve, and latex cast, x 4. 11, BC 12750, external mould of brachial valve, x 4. 12, BC

12751, internal mould of pedicle valve, x 4. 13, BC 12752, internal mould of pedicle valve, x 4. 14, BC 12753, internal mould of pedicle
valve, X 4. 15, BC 12754, external mould of brachial valve, x 4. 17, BC 12755, external mould of pedicle valve, x 4.

Fig. 16 Reuschella? sp. Kilbride. BC 12756, internal mould of pedicle valve, x 2.

Figs 18-20 Salopia sp. Fig. 18, Greenville-Moyne.BC 12757, internal mould of brachial valve, x 2. 19, 20, Kildare, Grange Hill Horizon 2.

BC 12758, internal mould of pedicle valve, and latex cast, x 2.

Figs 21-22 Porambonites sp. Kilbride. 21, BC 12759, internal mould of pedicle valve, x 2. 22, BC 12760, internalmould of pedicle valve, x 2.

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

160

69). Although closely similar, this form has an apparently
smooth bema and a greater frequency of accentuated ribs (at
least 7) than L. oepiki. They synonymized a population from
the Actonian of Shropshire which Hurst (1979a) had assigned
to L. oepiki, based only on internal moulds. The one figured
specimen of L. sp. (Hurst 1979a: 75, fig. 408), also from
Shropshire and synonymized by Harper & Owen (1984),
clearly shows a striated bema. Clearly, better preserved
material of all these forms would be desirable, especially
exteriors.

The inclusion of L. indentata in Bilobia by Cocks & Rong
(1989: 114) is considered erroneous, since the species does
not have the platform (sensu Cocks & Rong) near the
anterior margin, a feature obviously present in Bilobia
(Cocks & Rong 1989: 115, figs 70-71) but absent in Lep-
testiina. The bema is also more typically rounded and trans-
verse, aS in Leptestiina species, than the more complex
divided bema of Bilobia.

Leptestiina oepiki ampla subsp. nov. PI. 11, figs 7-17

NAME. Latin; ‘wide’.

DIAGNOSIS. Typical Leptestiina oepiki in all respects except
for the significantly more transverse bema than in the nomi-
nate subspecies.

HOLotyePe. BC 12748 (Pl. 11, figs 7, 8); paratypes BC
12749-55; Carrigadaggan.

MATERIAL AND LOCALITY. Carrigadaggan: 19 internal and 1
external moulds of pedicle valves; 2 internal and 2 external
moulds of brachial valves.

DISCUSSION. The new subspecies accords with the material
from Kilbride and Kildare, Grange Hill in all respects except
for one feature. The bema is more transverse, the mean
length being 46% of the width in two valves, as opposed to a
mean of 70% for 6 valves from Kilbride and 63% for 2 valves
from Grange Hill. It is difficult to assess theimportance of this
difference on such a small sample, but until the collection of
more topotype material proves otherwise the Carrigadaggan
population is assigned to the new subspecies L. oepiki ampla.

Family XENAMBONITIDAE Jones, 1928
Subfamily AEGIROMENINAE Havlicek, 1961
Genus CHONETOIDEA Jones, 1928

Chonetoidea abdita (Williams, in Whittington &
Williams 1955) Pl. 12, figs 10-16

1955 Sericoidea abdita Williams, in Whittington & Will-
iams: 418; pl. 39, figs 83-85.

MATERIAL AND LOCALITY. Greenville: 21 external and 14

PLATE 12

M. A. PARKES

internal moulds of pedicle valves, 32 external and 12 internal
moulds of brachial valves.

DESCRIPTION. Exterior. Concavo- or planoconvex, small
shells of transversely semicircular outline, maximum width at
hinge line. Cardinal angles acute to rectangular. Ventral
valve most convex near umbo in lateral profile, about 15% as
deep as long. Anterior profile rectimarginate, evenly convex,
occasionally strongly convex medially. Length 1s 58% of the
width (N = 53). Dorsal interarea hypercline and short,
ventral interarea apsacline. Ornament quite variable, com-
monly finely costellate, occasionally parvicostellate, with
about 10 ribs per mm, 2 mm anterior of the umbo.

Ventral interior. Very weakly impressed bilobed small
muscle field. Small, simple teeth project dorsilaterally and
are unsupported. Anterior margins of shell show feebly
developed ribs, sometimes extending posteriorly to mid-
valve. Interspaces and valve interior characterized by minute
pustules.

Dorsal interior. Short socket ridges ankylosed to small
cardinal process. Thin median septum extends to mid-length
of valve. Variable septule development, commonly 4, 5 or 6
septules arranged in an arc from the anterior end of the
median septum. Occasionally second anterior arc of septules
or very large circular pustules, third arc of coarse pustules
rarely developed. External ornament impressed on finely
pustulose interior.

Chonetoidea cf. abdita (Williams, in Whittington &
Williams 1955) Pl. 13, figs 14

MATERIAL AND LOCALITY. Kilbride: 6 internal and 2 exter-
nal moulds of brachial valves, 14 internal and 7 external
moulds of pedicle valves.

DESCRIPTION. Exterior. Small semicircular Chonetoidea,
widest at the hinge line. Concavoconvex profile, with maxi-
mum convexity at umbo, about 20% as deep as long. Recti-
marginate, with evenly convex anterior profile. Length about
64% of width. Hypercline, short dorsal interarea, apsacline
ventral interarea. Variable ornament, parvicostellate to
finely costellate, with occasional thickened ribs, with 15, 13
and 12 ribs per mm 2 mm anteriorly on 2, 1 and 1 valves.

Ventral interior. Small, weakly impressed muscle field.
Unsupported, short simple teeth. Interior shows relatively
strongly impressed ornament, particularly the accentuated
ribs of parvicostellate specimens.

Dorsal interior. Thin median septum extends over half the
valve length. Variable septule development in one or two
arcs, commonly 4 to 7 septules in the posterior are from the
anterior end of the median septum. External ornament is
impressed on interior, particularly the accentuated ribs of
parvicostellate specimens.

Discussion. The placing of Sericoidea in synonomy with

Figs 1-9 Bimuria cf. dyfiensis Lockley. Ballykale. 1, BC 12761, internal mould of pedicle valve, x 4. 2, 3, BC 12762, internal mould of pedicle
valve, and latex cast, x 48. 4, 5, BC 12763, internal mould of brachial valve, and latex cast, x 48. 6, BC 12764, dorsal view of conjoined
internal mould, x 48. 7, 8, BC 12765, external mould of brachial valve and part of pedicle valve, and latex cast showing interareas, x 48. 9,

BC 12766, external mould of brachial valve, x 48.

Figs 10-16 Chonetoidea abdita (Williams). Greenville. 10, BC 12767, internal mould of brachial valve, x 10. 11, BC 12768b, external mould
of pedicle valve, x 10. 12, BC 12769a, internal mould of brachial valve, x 4. 13, BC 12770b, external mould of brachial valve, x 4. 14, BC
12771, external mould of brachial valve, x 10. 15, BC 12772a, internal moulds of adjoined pedicle and brachial valves, x 4. 16, BC 12768a,

internal mould of pedicle valve, counterpart of Fig. 11, x 10.

;
‘BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND 161

M. A. PARKES

162

PS hy

e
! BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

_ MEASUREMENTS
_ Brachial valve internal moulds (Greenville):

|
'Variates X1 xD X24

Means 294 5-21 1-49
Sample size 12 12 9

| Variance-covariance 0-50 0:0 0-24
| Matrix 0-73 0-05

| 0-16

Brachial valve external moulds (Greenville):

Variates x1 X2
\Means 2:87 Seil7/
Sample size 32 32
Variance-covariance 0-34 0-20
Matrix 1-22

‘Pedicle valve internal moulds (Greenville):

Variates X1 X2 x4
‘Means 3-08 5:41 0-48
Sample size 14 14 9
/Variance-covariance 0-28 0:05 -0-02
Matrix 1:15 -0-01
| 0-01

Pedicle valve external moulds (Greenville):

Variates xl x2 x4

Means 2:99 5:29 0-52
Sample size 21 21 6
Variance-covariance 0:27 0:14 0-03
| Matrix 1:06 0-03
0-01

(Chonetoidea by Cocks & Rong (1989) completes a task
Suggested by many authors, including Hurst (1979a: 281) and
Young & Gibbons (1983), but not completed for lack of
sufficient material. The two described species are a clear
vindication of that decision, possessing characters variable
between those previously characteristic of both genera. These
include the external ornament (parvicostellate (Sericoidea) to
finely costellate (Chonetoidea)) and septule arrangement

PLATE 13

L Fig. 2, x 10.

mould of pedicle valve, counterpart of Fig. 7, x 2.

163

MEASUREMENTS
Brachial valve internal moulds (Kilbride):

Variates X1 x2 x24
Means 2-00 3-25 1-05
Sample size 6 6 2
Variance-covariance 0-12 0-17

Matrix 0-63

Pedicle valve internal moulds (Kilbride):

Variates Xl x2 x4
Means 2-19 3-39 0-44

Sample size 14 14 14
Variance-covariance 0-08 0-03 0-02
Matrix 0-29 0-01
0-02

Pedicle valve external moulds (Kilbride):

Variates X1 X2 x4
Means DAG 3-49 0:54
Sample size 7 7 7
Varianee-covariance 0-12 0-18 0-02
Matrix 0-55 0-05
0-01

(few septules in one arc (Sericoidea) to many septules and
more than one arc (Chonetoidea)). The interarea preserva-
tion is not good enough to discern whether canals are present,
as defined by Mitchell (1977: 93) in Chonetoidea.

Although the generic assignment is clearly to Chonetoidea,
the assignment of the two samples to Chonetoidea abdita and
Chonetoidea cf. abdita is made with some hesitation. Other
workers have found it difficult to distinguish species effec-
tively on the basis of shape or outline. The only commonly
variable characters found to be useful are the frequency of
costellae and the arrangement and number of septules or
sub-median septae in the lophophore platform of the brachial
valve. Comparisons with described species are hindered by
differences in numerical description. For example, older
descriptions note the number of costellae per mm at the
anterior margin irrespective of size, whilst later workers and
my Own counts were per mm at 2mm sagittally. Some recent
work has reverted to marginal counts, e.g. Harper (1989). In

Figs 1-4 Chonetoidea cf. abdita (Williams). Kilbride. 1, BC 12773, internal mould of pedicle valve, x 10. 2, BC 12774b, external mould of
brachial valve, x 10. 3, BC 12808, external mould of pedicle valve, x 10. 4, BC 12774a, internal mould of brachial valve, counterpart of

igs 5-8 Anisopleurella cf. multiseptata (Williams). Greenville-Moyne. 5, BC 12776a, internal mould of brachial valve, x 4. 6, BC 12776b,
external mould of brachial valve, counterpart of Fig. 5, x 4. 7, BC 12777b, external mould of pedicle valve, x 2. 8, BC 12777a, internal

figs 9-18 Sowerbyella sericea (J. de C. Sowerby). Figs 9-14,17, Kildare, Grange Hill Horizon 1. 9, BC 12778, internal mould of pedicle valve,

| x 4. 10, BC 12779, internal mould of pedicle valve, x 4. 11, BC 12780, internal mould of pedicle valve, x 4. 12, BC 12781, internal mould

| of pedicle valve, x 4. 13, BC 12782, internal mould of pedicle valve, x 4. 14, BC 12783, internal mould of brachial valve, x 4. 17, BC

| 12785, external mould of brachial valve and interareas, x 4. Figs 15, 16, 18, Kilbride. 15, 16, BC 12784, internal mould of brachial valve,
latex cast and mould, x 4. 18, BC 12786, external mould of brachial valve, x 4.

*igs 19-22 Ptychoglyptus sp. Kilbride. All incomplete external (?) moulds. 19, BC 12787, x 4. 20, BC 12788, x 4. 21, BC 12789, x 5. 22, BC

12790, x 4.

164

addition, the precision of the counts is slightly suspect, given
the very small size of specimens.

Notwithstanding the problem of definition of septules
versus coarse pustules (see Lockley, 1980: 215), the develop-
ment of second arcs and more septules is not dependent on
size, since smaller specimens can have as many or more than
medium or large specimens in the two samples. Although
Lockley (1980: 214, fig. 63) gives a useful table of the number
of septules in different size classes, it is not clear whether this
is total numbers or of one particular arc where more than one
is developed. In the Greenville sample, 4 out of 12 measured
brachial valves have second arcs developed with 2, 9, 12 and
13 septules present, and the third one has a third arc
developed with 11. In isolation, these would probably be
identified as Chonetoidea papillosa or Chonetoidea radiatula.
In the Kilbride species, 2 out of 6 had a second arc with 6 and
12 septules.

Neither species is like the type species Chonetoidea restricta
(Hadding) or Chonetoidea homolensis Havli¢ek (see Harper,
1989) as the median septum and septule arcs in these species
are contained within the posterior half of the shell. Although
contemporaneous, Chonetoidea sp. from the Gelli-grin For-
mation (Williams 1963) differs in having a platform devel-
oped by coalescence of strong septules. Percival (1979: 115)
described two species, Chonetoidea sejuncta and Chonetoidea
minor, which together with Chonetoidea virginica (Cooper,
1956) form a distinct species group. These have a thin median
septum extending up to mid-length, but the 2 to 4 pairs of
discrete small septules are positioned laterally in a row, not
arcuate as in the Greenville and Kilbride samples.

In summary, both the Leinster species are best assigned to
the Chonetoidea abdita Williams form, as emended by Lock-
ley (1980). The original species description was limited owing
to paucity of material. Williams has compared Lower Cara-
doc material from the Balclatchie Group (1962) of Girvan
and the Soudleyan Hagley Shales of Shropshire (1974) with
the Welsh form, and noted little difference. Although the
poorly preserved Kilbride form is here described separately,
as it is somewhat deeper and has a slightly greater frequency
of costellae than the Greenville sample, these differences are
probably not important. All the forms compared to the
species, including these samples, are probably displaying a
range of variation wider than that normally seen in the often
small samples available. The actual data for septule and
costellae numbers are given for comparison (Tables 21-22).

Table 21 Frequency of counts of costellae per mm at 2 mm
sagitally for Chonetoidea abdita from Greenville, Chonetoidea ct.
abdita from Kilbride, Chonetoidea cf. abdita from Shelva (data
from Williams, 1974) and at the anterolateral margins for
Chonetoidea aff. abdita from Girvan (data from Williams, 1962).

8) OOM OMIS a Aw Selomlotal
a} A I) 3) By il KO) C. abdita Greenville
il il 2 4 C. cf. abdita Kilbride
Sehr il 7 C. cf. abdita Shelve
3°97 & © 3B It Ae C. aff. abdita Girvan

|
M. A. PARKES |

Table 22 Distribution of various types of lophophore platform, |
with number of septules on either side of median septum (MS) in
columns for Chonetoidea abdita samples. Data for Chonetoidea
abdita from Greenville (left columns) and Chonetoidea cf. abdita
from Kilbride (central columns) based on posterior arc of larger
septules when more than one developed. Date for Chonetoidea
aff. abdita from Girvan (right columns) modified from Lockley
(1980).

Length MS 1 2 3 Total |
(mm) |
0-6-1-0 003 00 3
1-1-1-5 001 001 01 0 01 2
1-6-2-0 002 020 10 4 L216
2-1-2:5 002 210 ie 3) 312 Se
2-6-3-0 300 00 4 30 4
3-1-3-5 100 OS 30 35
3-6-4-0 20 0 20 0
004 005 630 62 14 WS 23}

Genus ANISOPLEURELLA Cooper, 1956

Anisopleurella cf. multiseptata (Williams, in
Whittington & Williams 1955) Pl. 13, figs 5-8)

MATERIAL AND LOCALITY. Greenville-Moyne: 8 internal and,
2 external moulds of pedicle valves, 2 internal and 5 external
moulds of brachial valves.

DISCUSSION. The sparse material, especially brachial valve
interiors, is inadequately preserved
deformed, making a specific determination difficult. Brench-

ley et al. (1977) recorded Anisopleurella aff. multiseptata’
Williams from this locality, but although similar to that —
species, the sample is apparently more transverse. Lockley

(1980), however, compared a single pedicle valve from the

Gelli-grin Formation of Bala to the species A. multiseptata

which appears similar to the material from Greenville-

Moyne. The exteriors are poor but show the essentially

smooth exterior ornamented by a few widely-spaced primary

costae, and Brenchley er al. (1977) are followed in compari-

son of this species to A. multiseptata Williams.

Family SOWERBYELLIDAE Opik, 1930
Subfamily SOWERBYELLINAE Opik, 1930
Genus and subgenus SOWERBYELLA (SOWERBYELLA)
Jones, 1928

Sowerbyella sericea (J. de C. Sowerby, 1839)

Pl. 13, figs 9-18)
Leptaena sericea J. de C. Sowerby, in Murchison‘
636; pl. 19, fig. 1.
Sowerbyella sericea (J. de C. Sowerby) Jones: 414:
pl. 21, figs 1-4.
Sowerbyella sericea (J. de C. Sowerby); Williams:
430-432; pl. 11, figs 1-9.
Sowerbyella sericea (J. de C. Sowerby); Bretsky:
85-87; pl. 12, figs 3-6; pl. 13, figs 1-4.
Sowerbyella sericea (J. de C. Sowerby); Williams:
134-135; pl. 24, figs 11-14, 16.
Sowerbyella sericea (J. de C. Sowerby); Cocks: 98.

1839

1928

1963

1970

cf.1974

1978

and moderately) —

|
/

_ BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

1979a Sowerbyella sericea (J. de C. Sowerby); Hurst: 278;
figs 412-432.

' MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon
_1: 3 external and 15 internal moulds of pedicle valves; 7
' external and 4 internal moulds of brachial valves. Kildare,
Grange Hill, Horizon 2: 2 internal moulds of pedicle valves.
Kilbride: 9 external and 17 internal moulds of pedicle valves,
_7 external and 2 internal moulds of brachial valves. Carri-
gadaggan: 1| internal mould of a pedicle valve; 4 external and
| 2 internal moulds of brachial valves.
DESCRIPTION. Exterior. Semicircular outline, cardinal angles
acute in smaller specimens, becoming rectangular in older
stages. Concavo-convex shells, with median fold occasionally
developed in pedicle valve, and concave flanks on pedicle
valve. Length about 55-60% of the width, and depth about
one third of the length in the pedicle valve. Radial ornamen-
tation unequally parvicostellate, but quite variable, with 6-10
costellae per mm, 5 mm anteromedially of the umbo, segre-
gated into sectors about 1 mm wide if not finely costellate.
| Occasionally a few pairs of rugae developed in posterolateral
areas. Dorsal interarea very short, flat and catacline, ventral
‘interarea curved and apsacline.

Ventral interior. Bilobed, divergent diductor muscle scars
_about 0-7 as long as wide, extending anteriorly for about
one-third of valve length. Adductor scars separated posteri-
|orly by thin median septum extending for 15-20% of valve
length before bifurcating. Diductors widely separated anteri-
orly and split by divergent vascula media bounded by lateral
ridges. Lemniscate pallial sinus pattern, with papillose ante-
rior surface. Small teeth with obsolescent dental plates in
adults.
| Dorsal interior: Undercut cardinal process fused with
widely divergent socket ridges. The bases of the socket ridges
jare continuous with two low ridges that rise in height anteri-
orly to become prominent sub-median septa extending about
two-thirds of valve length. Sub-median septa diverge slightly,
so that anterior separation is about one-third of their length.
Low, small median septum and thinner, low, radial ridges
,across bema occasionally developed. Lemniscate pallial sinus
|pattern, and papillose interior anterior of the bema.

Discussion. Although differing slightly in some propor-
tions, from each other and from the type material, the
differences are not significant, given the variability known in
stocks of Sowerbyella (Cocks & Rong 1989: 139) and do not
justify erection of a subspecies. A few of the measured
specimens are also slightly distorted so the reliability of the
Statistics is low, but is presented as a general assessment of
the morphology. Material from the Grange Hill, Carrigadag-
gan and Kilbride populations is all referred to the type species

5 sericea (J. de C. Sowerby). The type species was revised by
Williams (1963), and Hurst (1979a) figured a large sample of
this species from the Woolstonian of south Shropshire to
illustrate the variability within the species. The number of
dorsal valves is small, but plots of the first four eigenvectors
(Fig. 19) from a principal component analysis of pedicle
valves from Kildare, Grange Hill and Kilbride shows no
differences between the samples, except for overall size
where vector 1 shows some differentiation. The Kilbride
Fanon has a smaller mean size.

|
|

165

MEASUREMENTS
Brachial valve external moulds (Kilbride);

Variates X1 X2 x4
Means 6:83 1:21 1-35,
Sample size if 7 6
Variance-covariance 2:08 4-46 0-27
Matrix DeSwil — (OeSi7/
0-26

Pedicle valve internal moulds (Kilbride):

Variates XT XOX XO Xd XO4
Means 5:51 9-42 1-83 1-43 2-10 0-77
Sample size Wy i 1 6 9

4-06 4-79 1-11 1-08 1:74 0-78
0-70 0-23 0-43 0-28
0230-27019

0-49 0-24

Variance-covariance
Matrix

Brachial valve internal moulds (Kildare, Horizon 1):

Variates WIL KDB) DP). 9-73}
Means 9-38 1:78 2-85 6:08 1-95
Sample size 4 4 4 4 4

Variance 1:17 6-09 0-37 0-32 0-01

Brachial valve external moulds (Kildare, Horizon 1):

Variates Xl X2 X4
Means 7-96 1-43 1-38
Sample size 7 7 6
Variance-covariance 13-43 17-64 1-55
Matrix 32-97 3-44
0-57

Pedicle valve internal moulds (Kildare, Horizon 1):

Variates XC | XOX XO xd OR X24:
Means 9-47 15:2 3-53 4-06 5-76 2-20
Sample size sy 15 Ss le Aas

3-61 4:97 0-69 1-88 2:33 0-86
16-24 2:79 2-62 5-77 1-54
1-48 0:52 1-27 0-29

1:24 1-38 0-51

2°62 0-77

Variance-covariance
Matrix

Pedicle valve external moulds (Kildare, Horizon 1):

Variatesa Xl X2 x4
Means 9-47 16-0 3:27
Sample size 3 3 3
Variance 1-04 2:82 0-22

166

Subfamily PTYCHOGLYPTINAE Cooper, 1956
Genus PTYCHOGLYPTUS Willard, 1928

Pins stipseo 27

MATERIAL AND LOCALITY. Kilbride: All material incom-
plete; one internal and one external moulds of a pedicle
valve, one internal and one external moulds of a brachial
valve, and 8 indeterminate moulds.

Ptychoglyptus sp.

DESCRIPTION. Characteristic Ptychoglyptus rugae with a
wavelength of 2 per mm, separated into discontinuous arcu-
ate chevron-like pattern by strong radial costellae. Whole of
surface, including rugae, sculpted with fine radial lines.
Straight hinge line and slight rafinesquinoid convexity in
pedicle valve. The dorsal interior is inadequately known.

MEASUREMENT. One specimen shows most of half a hinge
line and gives a minimum hinge width of 21 mm.

DISCUSSION. The material is too fragmentary to justify a
specific determination. It appears to be unlike P. ? kindlei
Cooper (1956) which has a geniculate anterior, and also
unlike P. ? matura Cooper (1956) which is non-rugose in the
anterior half. It is comparable to P. virginiensis Willard
(1928) in most respects except for the rugose sculpture. As
Williams (1962) noted, this species has rounded crests to the
rugae and is generally symmetrical or asymmetrical with
steeper posterior slopes. Williams (1962) described three
specimens of P. cf. valdari from Balclatchie, near Girvan in
Scotland, with sharply asymmetrical rugae, and three with
rugae overfolded in an anterior direction. The material from
Kilbride most closely resembles this species since the crests
are sharp and asymmetrical. The posterior slope is steeper
but in some rugae there is an anteriorly deflected crest,
creating an overfolded appearance, but it is variably devel-
oped within single specimens.

Superfamily STROPHOMENOIDEA King, 1846
Family STROPHOMENIDAE King, 1846
Subfamily STROPHOMENINAE King, 1846
Genus STROPHOMENA Rafinesque, 1825

Pl. 14, figs 13, 6

MATERIAL AND LOCALITY. Kildare, Grange Hill, Horizon 1:
2 external and 1 internal moulds of a pedicle valve.

Strophomena ? sp

DESCRIPTION. Gently, evenly convex pedicle valve with very
shallow sulcus. Interarea high, apsacline about 14% as long
as the valve. Simple triangular teeth supported by short
divergent dental plates. Delthyrium almost completely closed
by convex pseudodeltidium. Faint transverse grooves and
ridges on the interarea near the hinge line. Interior shows

PLATE 14

Figs 1-3, 6 Strophomena? sp. Single specimen from Kildare, Grange Hill Horizon 1. 1, 2, BC 12791b, external mould of pedicle valve, and |
latex cast, X 2. 3, 6, BC 12791a, counterpart internal mould of pedicle valve, and latex cast, x 2.

Fig. 7 Kjerulfina? sp. Kildare, Grange Hill Horizon 1. BC 12792, internal mould of pedicle valve, x 2.

Figs 4-5, 8-10, 14-17 Rafinesquina sp. Kildare, Grange Hill House Cottage. 4, 5, BC 12793, internal mould of brachial valve, latex cast and
mould, x 2. 8, BC 12794, internal mould of pedicle valve, x 175. 9, BC 12795b, external mould of pedicle valve, x 2. 10, 14, BC 12795a,
internal mould of pedicle valve, counterpart of Fig. 9, latex cast and mould, x 2. 15, BC 12796b, external mould of brachial valve, x 2. 16,
17, BC 12796a, internal mould of brachial valve, counterpart of Fig. 15, latex cast and mould, x 2.

Figs 11-13 Hedstroemina sp. Kildare, Grange Hill Horizon 1. 11, BC 12797a, internal mould of pedicle valve, x 2. 12, BC 12798, internal
mould of pedicle valve, x 2. 13, BC 12799, internal mould of pedicle valve, x 2.

M. A. PARKES

impression of ornament throughout, especially at margins.
External ornament finely costellate with about 6 ribs per mm
at 5 mm growth stage. Strong concentric, accentuated growth
lines on exterior of shell.

MEASUREMENTS. BC 12791a & b: X1 = 28 mm, X2 = 21
mm, X4 = 3 mm, X6 = 38 mm; X1/X2 = 0-75, X4/X1 = 0-11,
X6/X1 = 0-14 (PI. 14, figs 1-3, 6).

DiscussION. Although the unequally parvicostellate orna-
ment is lacking, the single valve part and counterpart moulds
are questionably assigned to Strophomena, as the general
morphology corresponds to the genus in other aspects. How-
ever it is 75% as wide as long, unlike species from Wales that _
are wider than long; there is no apparent deformation of the
specimen.

Genus KJERULFINA Bancroft, 1929

Pl. 14, fig. 7

MATERIAL AND LOCALITY. Kildare, Grange Hill, Horizon 1:
a single internal mould of pedicle valve, incomplete.

MEASUREMENTS. BC 12792: X1 = 15 mm, X4 = 33 mm (PI.
14, fig. 7).

DISCUSSION. This incomplete mould has an initially gently
convex profile, but deflects abruptly in a ventral direction
before geniculating sharply in a dorsal direction. It is moder-
ately wrinkled by rugae posterolaterally and is finely orna-
mented by costellae impressions. The dental plates are
divergent, short and curving. The specimen exhibits similari-
ties to Kjerulfina, one species of which, K. broeggeri, has
similar sporadic dorsally directed geniculation. Further mate-
rial is required to reach a better identification.

Kjerulfina ? sp.

Subfamily RAFINESQUININAE Schuchert 1893
Genus RAFINESQUINA Hall & Clarke 1892

Rafinesquina sp. Pl. 14, figs 4-5, 8-10, 14-17 |

MATERIAL AND LOCALITY. Kildare, Grange Hill House Cot-_
tage: 5 internal and 5 external moulds of pedicle valves, 8 |
internal and 4 external moulds of brachial valves, 4 indeter- |
minate external moulds and 3 indeterminate internal moulds, |
mostly fragmentary.

DISCUSSION. The best specimens of this genus are figured to

show the typical form of the sample, but it is not described |
pending collection of more material to assess the morphologi- |
cal variation. It shows typical concavo-convex form, and |
unequally parvicostellate ornamentation. However, there are
few modern descriptions of the genus from Britain or Ireland

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND

167

168

2.9

1.7 229
SOWPV2

1.7

2.9
SOWV2

A0),7/
V-2

0.5

M. A. PARKES

and few well-defined species. Lamont (1953) claimed the
record by Reynolds & Gardiner (1896) of Strophomena
expansa J. de C. Sowerby should be Rafinesquina concentrica
(Portlock) var. However, specimens collected by Lamont
(NMING: F 18662, F18663, F18599, F18611) from this locality
are labelled as ‘R. eirenach Lamont’. No publication of this
species name is known, and the material is kept in open
nomenclature, pending further collection and analysis.

Family STROPHOMENIDAE King, 1846
Genus HEDSTROEMINA Bancroft, 1929

Hedstroemina sp.

MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon
1: 3 internal and 3 external moulds of pedicle valves; 3
external moulds of brachial valves. Kildare, Grange Hill,
Horizon 2: 7 internal moulds of pedicle valves.

DISCUSSION. The restricted sample makes identification
problematical, given the wide variation within the genus and
the closely related Kjaerina. As also noted by Williams (1963:

460), in a very small Bala sample the lack of a strong median -

rib invites allocation to Hedstroemina rather than Kjaerina.
Hurst (1979a: 288) notes four characteristics in Upper Cara-

doc populations which could consistently be used to separate |

them, one being the development of weak rugae which is also
seen in this material.

Family LEPTAENIDAE Cooper, 1956
Genus LEPTAENA Dalman, 1828

Leptaena sp. Pl. 15, figs 19

MATERIAL AND LOCALITIES. Kildare, Grange Hill, Horizon
1: 6 internal and 3 external moulds of pedicle valves; 5
external moulds of brachial valves; 6 indeterminate exterior
fragments and 1 conjoined internal mould. Kildare, Grange
Hill, Horizon 2: 1 internal and 1 external moulds of pedicle
valves; | internal and 2 external moulds of brachial valves, all
incomplete.

DISCUSSION. The small fragmentary sample is insufficient for
statistical comparison with other known Leptaena species,
and is left under open nomenclature pending better material
to assess the variability. Inspection of the ventral muscle
field, however, suggests it may be conspecific with L. ventri-
cosa Williams from the Longvillian Gelli-grin Group of Bala.

Fig. 19 Principal component analysis of Sowerbyella sericea
(pedicle valve internal moulds) from Kildare, Grange Hill
Horizon 1 (W), Kilbride (*) and Carrigadaggan (0). A, Plot of
vector 1 against vector 2; B, Plot of vector 3 against vector 4; C,
Plot of vector 2 against vector 3.

Pl. 14, figs 11-13 |

BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND
|

Family STROPHEODONTIDAE Caster, 1939

| Subfamily STROPHEODONTINAE Caster, 1939

_ Genus HIBERNODONTA Harper & Mitchell, in Harper et
. al. 1985

Hibernodonta ? sp. Pl. 15, figs 8-10
MATERIAL AND LOCALITY. Kildare, Grange Hill House Cot-
tage: 3 internal and 1 external mould of brachial valves, 2
external moulds of pedicle valves and 3 indeterminate exter-
nal moulds.

Discussion. The genus and species H. praeco were first
described from the Clashford House Formation at Herbert-
'stown, Co. Meath by Harper & Mitchell (in Harper et al.
1985). As an early stropheodontid, the small rafinesquinid-
like valves have denticulate hinge lines. The specimens here
sare tentatively assigned to Hibernodonta, since there is an
‘appearance of denticulation on the hinge line, as well as
definite denticulate teeth. However, the material is poorly
preserved and only three valves have the hinge area present,
/although the size of the sample is larger than that of the type
species. There is no thickened median rib. Until better
preserved specimens are collected the material can only be
-questionably assigned to Hibernodonta; it may equally be a
strophomenoid species which developed denticulate teeth
}and sockets, other cases of which are discussed by Harper et
val. (1985).

(PLATE 15

| House Cottage, BC 12806a, internal mould of pedicle valve, x 2.

169

Superfamily PORAMBONITOIDEA Davidson, 1853
Family PORAMBONITIDAE Davidson, 1853
Genus PORAMBONITES Pander, 1830

Porambonites sp. Pl. 11, figs 21-22

MATERIAL AND LOCALITY. Kilbride: 2 internal moulds of
pedicle valves.

DESCRIPTION. Ventral interior. Large valves of subtriangular
outline, with strong convexity posteriorly. Shallow sulcus
originating about mid-length. Faint growth line impressions
near anterior margin. Strong teeth supported by high, thin
parallel dental plates extending anteriorly for nearly half of
length. (Exteriors are unknown.)

MEASUREMENTS. BC 12670: (X1) length = 23 mm, (X2)
width = 18-4 mm, (X3) position of maximum width = 16-5
mm, (X4) depth = 7-4 mm; length of dental plates = 11-4 mm
(PI. 11, fig. 22).

DISCUSSION. The material is inadequate for specific assign-
ment, but may be conspecific with the poorly known Poram-
bonites filosus M’Coy from nearby Knockmahon, Co.
Waterford. Porambonites is a common genus in the TLF
(Carlisle 1979) and is currently under revision by Parkes &
Harper. Significant differences, however, appear between
this species and the older one in size, outline and internal
morphology.

\Figs 1-7 Leptaena sp. Figs 1-4, 6-7, Kildare, Grange Hill Horizon 1. 1, BC 12800, external mould of brachial valve, x 2. 2, BC 12801,
| internal mould of pedicle valve, x 2. 3, BC 12802, external mould of brachial valve, x 2. 4, BC 12803, internal mould of pedicle valve, x
2. 6, BC 12804, external mould of brachial valve, x 2. 7, BC 12805, internal mould of pedicle valve, x 2. Fig. 5, Kildare, Grange Hill

Figs 8-10 Hibernodonta? sp. Single specimen from Kildare, Grange Hill House Cottage. 8, 10, BC 12807a, internal mould of brachial valve,
and latex cast, x 4. 9, BC 12807b, counterpart external mould of brachial valve, x 4.

170

M. A. PARKES

f)
| BRACHIOPODS OF DUNCANNON GROUP OF SE IRELAND
ACKNOWLEDGEMENTS. I wish to thank David Harper for constant
) enthusiastic direction and support; all the University College Galway
'(U.C.G.) Geology Department staff for assistance and support, and
Lucy Kellehan for her patient help. Many thanks to my parents for
their continuous help and encouragement, and also to Carol Gray for
‘her support. I gratefully acknowledge a U.C.G. Postgraduate Fellow-
ship (1985-86) and an Anglo-Irish Agreement Postgraduate Scholar-
ship. An Irish Geological Association grant and European Union of
Geosciences Travel Grant assisted presentations of some of this
work. Thanks also to the many farmers who have allowed access and
camping on their lands; in particular Liam and Emily Murphy of
|Raheen, Tim Murphy of Wilton Mills, Hilary Powers of Kilbride and
Tom Foley of Carrigadaggan. Much help and advice has been freely
‘given by many people in the geology departments of Trinity College

Dublin (T.C.D.), University College Dublin, University College
Cork, the G.S.I. and elsewhere, but particular thanks go to J.
‘Jackson (deceased), N. Monaghan, S. Morris, F.C. Murphy, A.W.

Owen, C.R.C. Paul, P. Shannon and D. Tietzsch-Tyler, as well as J.
Wilson (Ulster Museum). The referee’s comments considerably
improved the work. Also, C.H. Holland permitted use of the
‘facilities in T.C.D. to complete this Bulletin. Thanks to Conall
/MacNiocaill for photographic help, and I especially thank Michelle
Davern.

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Journal of Science, New Haven, (5) 20: 265-288, pls 1-3.

—— 1932. Brachiopod genera of the suborders Orthoidea and Pentam-
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1-270.

—— & Le Vene, C.M. 1929. Brachiopoda (Generum et genotyporum index et
bibliographia). Fossilium Catalogus, Berlin, 1 Animalia (42). 140 pp.

Sedgwick, A. & M’Coy, F. 1851-55. A synopsis of the classification of the
British Palaeozoic rocks, with a systematic description of the British Palaeo-
zoic fossils in the geological museum of the University of Cambridge. xcviii +
661 pp., 25 pls. London and Cambridge.

Shannon, P.M. 1977. Early Caledonian dolerites and intrusion-related struc-
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1: 25-31.

—— 1978. The stratigraphy and sedimentology of the Lower Palaeozoic rocks
of south-east County Wexford. Proceedings of the Royal Irish Academy,
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— 1979a. The petrology of the Ordovician volcanic rocks of County
Wexford, Ireland. Journal of Earth Sciences, Royal Dublin Society, 2: 41-59.

— 1979b. The tectonic evolution of the lower Palaeozoic rocks of extreme SE
Ireland. /n: Harris, A.L. et al. (eds), The Caledonides of the British Isles —
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1980. Depositional mechanisms and environments in the Lower Palaeo-
zoic clastic sediments of southeast Ireland. Journal of Earth Sciences, Royal
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Sheehan, P.M. 1987. Late Ordovician (Ashgillian) brachiopods from the region
of the Sambre and Meuse Rivers, Belgium. Bulletin de l'Institut Royal des
Sciences Naturelles de Belgique, Brussels, (Sci. Terr.) 57: 5-81.

Smyth, L.B. 1939. The geology of south-east Ireland, together with parts of
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Sowerby, J. de C. 1839. In Murchison, R.I.,
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Spjeldnaes, N. 1957. The Middle Ordovician of the Oslo Region, Norway. 8.
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Oslo, 37: 1-214.

Stillman, C.J. 1976. A Field Guide to the Ordovician Volcanic Rocks of
South-East County Waterford. Geology Department, Trinity College
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Temple, J.T. 1970. The Lower Llandovery brachiopods and trilobites from
Fridd Mathrafal, near Meifod, Montgomeryshire. Monograph of the Palae-
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1987. Early Llandovery brachiopods of Wales. Monograph of the Palaeon-
tographical Society, London. 137 pp., 15pls.

Tietzsch-Tyler, D. 1989. The Lower Palaeozoic geology of S.E. Ireland — a
revaluation. Irish Association for Economic Geology, Annual Review, Dub-
lin, 1989: 112-119.

Ulrich, E.O. & Cooper, G.A. 1936. New Silurian Brachiopoda of the family
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1992. New Caradoc brachiopods from the Iberian Chains (northeastern

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1963. The Caradocian Brachiopod Faunas of the Bala District, Merione-

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8 (7): 327-471.

1974. Ordovician Brachiopoda from the Shelve District, Shropshire.

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ment 11: 1-163.

M. A. PARKES

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makeady Limestone, Co. Mayo, Ireland. Bulletin of the British Museum |
(Natural History), London, Geology 38 (4): 183-269. )

— Lockley, M.G. & Hurst, J.M. 1981. Benthic palaeocommunities repre- |
sented in the Ffairfach Group and coeval Ordovician successions of Wales.
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— et al: 1965. Treatise on Invertebrate Palaeontology, H. Brachiopoda. 927 |
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Williams, J.S. 1943. Stratigraphy and fauna of the Louisiana Limestone of ©
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Woodward, S.P. 1852. A manual of the Mollusca; or rudimentary Treatise of |
recent and fossil shells. xvi + 486 pp., 24 pls. London.

Wright, A.D. 1963. The Fauna of the Portrane Limestone; part 1, the |

Inarticulate Brachiopods. Bulletin of the British Museum (Natural History), |
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— 1970. The stratigraphic distribution of the Ordovician inarticulate brachio- |
pod Orthisocrania divaricata (M’Coy) in the British Isles. Geological Maga- |
zine, Cambridge, 107: 97-103. :
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Sheffield Geological Society, 8.2: 95-104.

Bulletin of The Natural History Museum
Geology Series

Earlier Geoiogy Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete

| backlist is not shown, this may also be obtained from the same address.

. Volume 31
‘No. 1

| No. 2

No. 3

No. 4

| Volume 32
‘No. 1

No. 4

‘Volume 33
No. 1

No. 2

Foraminifera of the Togopi Formation, eastern Sabah,
Malaysia. J.E. Whittaker & R.L. Hodgkinson. 1979. Pp.
1-120, 10 plates, 71 figs. £14.00

Cretaceous faunas from Zululand and Natal, South
Africa. The ammonite family Gaudryceratidae. W.J.
Kennedy & H.C. Klinger. 1979. Pp. 121-173. £6.25

Benthic community organization in the Ludlow Series of
the Welsh Borderland. R. Watkins. 1979. Pp.

175-279. £12.25
The ammonites of the English Chalk Rock (Upper
Turonian). C.W. Wright. 1979. Pp. 281-330. £6.50
Miscellanea

Observations on Cycloclypeus (Cycloclypeus) Carpenter
and Cycloclypeus (Katacycloclypeus) Tan
(Foraminiferida). C.G. Adams & P. Fame. 23 figs.

The provenance of Sivapithecus africanus. P.J. Andrews
& T.I. Molleson. 3 tables.

A silicified brachiopod fauna from the Silurian of Iran.
L.R.M. Cocks. 41 figs.

Two new condylarths (Mammalia) from the early Eocene
of southern England. J.J. Hooker. 15 figs, 3 tables.
Miocene sharks’ teeth from Ecuador. A.E. Longbottom.
26 figs, 3 tables.

A new fossil terrestrial isopod with implications for the
East African Miocene land form. S.F. Morris. 12 figs.

A re-evaluation of the fossil human calvaria from Singa,
Sudan. C.B. Stringer. 2 figs, 1 table.

New species of Protorthoptera and Protodonata (Insecta)
from the Upper Carboniferous of Britain, with a
comment on the origin of wings. P.E.S. Whalley. 5 figs.
1979. Pp. 1-90. £10.50

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

The Ordovician trilobite faunas of the
Builth-Llandrindod Inlier, central Wales. Part III. C.P.
Hughes. 1979. Pp. 109-181, 177 figs. £10.00

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

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. £10.00
Map available separately £1.00

Miscellanea

A new, possibly algal, microproblematicum from the
Lower Carboniferous of England. G.F. Elliott, 8 Figs.
Acanthopleurella Groom 1902: origin and life-habits of a
miniature trilobite. R.A. Fortey & A.W.A. Rushton. 21
figs.

Pleistocene bird remains from Tornewton Cave and the
Brixham Windmill Cave in south Devon. C.J.O.
Harrison. | fig.

The succession of Hyracotherium (Perissodactyla,
Mammalia) in the English early Eocene. J.J. Hooker, 6
figs.

Salenia trisuranalis sp. nov. (Echinoidea) from the
Eocene (London Clay) of Essex, and notes on its
phylogeny. D.N. Lewis & R.P.S. Jefferies. 5 figs.
Tertiary and Cretaceous brachiopods from Seymour,
Cockburn and James Ross Islands, Antarctica. E.F.
Owen. 33 figs.

Revision of the rugose coral Diphyllum concinnum
Lonsdale, 1845, and historical remarks on Murchison’s
Russian coral collection. B.R. Rosen & R.F. Wise. 3
figs.

Neuroptera (Insecta) in amber from the Lower
Cretaceous of Lebanon. P.E.S. Whalley. 12 figs. 1980.

Pp. 71-164. £12.00
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
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
No. 5 The Yorkshire Jurassic fern Phlebopteris braunii
(Goeppert) and its reference to Matonia R.Br. T.M.
Harris. 1980. Pp. 295-311, 2 figs. £2.75
Volume 34
No. 1 Relative dating of the fossil hominids of Europe. K.P.
Oakley. 1980. Pp. 1-63, 6 figs, 17 tables. £8.00
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

No. 3 Ashgill Brachiopoda from the Glyn Ceiriog District,
north Wales. N. Hiller. 1980. Pp. 109-216, 408
figs. £14.75

No. 4 Miscellanea
Type specimens of some Upper Palaeozoic Athyridide
brachiopods. C.H.C. Brunton. 31 figs.
Two new British Cretaceous Epitoniidae (Gastropoda):
evidence for ev olution of shell morphology. R.J.
Cleevely. 14 figs, 1 table.
Revision of the microproblematicum Prethocoprolithus
Elliott, 1962. G.F. Elliott. 4 figs.
Basilicus tyrannus (Murchison) and the glabellar
structure of asaphid trilobites. R.A. Fortey. 12 figs.
A new Lower Ordovician bivalve family, the Thoraltidae
(? Nuculoidea), interpreted as actinodont deposit
feeders. N.J. Morris. 7 figs.
Cretaceous brachiopods from northern Zululand. E.F.
Owen. 13 figs.
Tupus diluculum sp. nov. (Protodonata), a giant
dragonfly from the Upper Carboniferous of Britain.
P.E.S. Whalley. 1 fig.
Revision of Plummerita Br6nniman (Foraminiferida) and
a new Maastrichtian species from Ecuador. J.E.

Whittaker. 34 figs. 1980. Pp. 217-297. £11.00
Volume 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

No. 2 The fossil alga Girvanella Nicholson & Etheridge.
H.M.C. Danielli. 1981. Pp. 79-107, 8 figs, 3 tables. £4.20

No. 4

Volume 36

No. 1

No. 3

No. 4

Volume 37

No. 1

Centenary miscellanea

Reassessment of the Ordovician brachiopods from the
Budleigh Salterton Pebble Bed, Devon. L.R.M. Cocks &
M.G. Lockley. 35 figs.

Felix Oswald’s Turkish Algae. G.F. Elliott. 3 figs.

J.A. Moy-Thomas and his association with the British
Museum (Natural History). P.L. Forey & B.G.
Gardiner. 3 figs.

Burials, bodies and beheadings in Romano-British and
Anglo-Saxon cemeteries. M. Harman, T.I. Molleson &
J.L. Price. 5 figs, 7 tables, VI appendices.

The Jurassic irregular echinoid Nucleolites clunicularis
(Smith). D.N. Lewis & H.G. Owen. 4 figs.

Phanerotinus cristatus (Phillips) and the nature of
euomphalacean gastropods. N.J. Morris & R.J. Cleevely.
12 figs.

Agassiz, Darwin, Huxley, and the fossil record of teleost
fishes. C. Patterson. 1 fig.

The Neanderthal problem and the prospects for direct
dating of Neanderthal remains. C.B. Stringer & R.
Burleigh. 2 figs, 1 table.

Hippoporidra edax (Busk 1859) and a revision of some
fossil and living Hippoporidra (Bryozoa). P.D. Taylor &
P.L. Cook. 6 figs. 1981. Pp. 109-252. £20.00

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

Middle Cambrian trilobites from the Sosink Formation,
Derik-Mardin district, south-eastern Turkey. W.T.
Dean. 1982. Pp. 1-41, 68 figs.

Miscellanea

British Dinantian (Lower Carboniferous) terebratulid
brachiopods. C.H.C. Brunton. 20 figs.

New microfossil records in time and space. G.F. Elliott.
6 figs.

The Ordovician trilobite Neseuretus from Saudi Arabia,
and the palaeogeography of the Neseuretus fauna related
to Gondwanaland in the earlier Ordovician. R.A. Fortey
& S.F. Morris. 10 figs.

Archaeocidaris whatleyensis sp. nov. (Echinoidea) from
the Carboniferous Limestone of Somerset and notes on
echinoid phylogeny. D.N. Lewis & P.C. Ensom. 23 figs.
A possible non-calcified dasycladalean alga from the
Carboniferous of England. G.F. Elliott. 1 fig.
Nanjinoporella, a new Permian dasyclad (calcareous
alga) from Nanjing, China. X. Mu & G.F. Elliott. 6 figs,
1 table.

Toarcian bryozoans from Belchite in north-east Spain.
P.D. Taylor & L. Sequeiros. 10 figs, 2 tables.

Additional fossil plants from the Drybrook Sandstone,
Forest of Dean, Gloucestershire. B.A. Thomas & H.M.
Purdy. 14 figs, 1 table.

Bintoniella brodiei Handlirsch (Orthoptera) from the
Lower Lias of the English Channel, with a review of
British bintoniellid fossils. P.E.S. Whalley. 7 figs.
Uraloporella Korde from the Lower Carboniferous of
South Wales. V.P. Wright. 3 figs. 1982. Pp.

43-155. £19.80

The Ordovician Graptolites of Spitsbergen. R.A. Cooper
& R.A. Fortey. 1982. Pp. 157-302, 6 plates, 83 figs, 2
tables. £20.50

Campanian and Mastrichtian sphenodiscid ammonites
from southern Nigeria. P,M.P. Zaborski. 1982. Pp.
303-332, 36 figs.

£5.80

£4.00

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

No.

Volume 38
No.

No.

Volume 39
No.

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

Miscellanea }
Glossopteris anatolica Sp. nov. from uppermost Permian
strata in south-east Turkey. S. Archangelsky & R.H.
Wagner. 14 figs. }
The crocodilian Theriosuchus Owen, 1879 in the

Wealden of England. E. Buffetaut. 1 fig.
A new conifer species from the Wealden beds of |
Féron-Glageon, France. H.L. Fisher & J. Watson. 10 '
figs. ]
Late Permian plants including Charophytes from the
Khuff formation of Saudi Arabia. C.R. Hill & A.A.
El-Khayal. 18 figs.
British Carboniferous Edrioasteroidea (Echinodermata). —
A.B. Smith. 52 figs.

A survey of recent and fossil Cicadas (Insecta,
Hemiptera-Homoptera) in Britain. P.E.S. Whalley. 11 figs. —
The Cephalaspids from the Dittonian section at Cwm Mill,
near Abergavenny, Gwent. E.I. White & H.A. Toombs. 20 |
figs. 1983. Pp. 79-171. £13.50

The relationships of the palaeoniscid fishes, a review
based on new specimens of Mimia and Moythomasia
from the Upper Devonian of Western Australia. B.G.
Gardiner. 1984. Pp. 173-428. 145 figs. 4 plates. 0 565
00967 2. £39.00

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 |

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 |

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

|
|
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 |
|

Miscellanea

Growth and shell shape in Productacean Brachiopods.
C.H.C. Brunton.

Palaeosiphonium a problematic Jurassic alga. G.F.
Elliott.

Upper Ordovician brachiopods and trilobites from the
Clashford House Formation, near Herbertstown, Co.
Meath, Ireland. D.A.T. Harper, W.I. Mitchell, A.W.
Owen & M. Romano.

Preliminary description of Lower Devonian Osteostraci
from Podolia (Ukrainian S.S.R.). P. Janvier.

Hipparion sp. (Equidae, Perissodactyla) from Diavata
(Thessaloniki, northern Greece). G.D. Koufos.
Preparation and further study of the Singa skull from
Sudan. C.B. Stringer, L. Cornish & P. Stuart-Macadam.
Carboniferous and Permian species of the cyclostome
bryozoan Corynotrypa Bassler, 1911. P.D. Taylor.
Redescription of Eurycephalochelys, a trionychid turtle
from the Lower Eocene of England. C.A. Walker &
R.T.J. Moody.

Fossil insects from the Lithographic Limestone of
Montsech (late Jurassic-early Cretaceous), Lérida
Province, Spain. P.E.S. Whalley & E.A. Jarzembowski. |
1985. Pp. 271-412, 162 figs. 0 565 07004 5. £24.00 |

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

Cenomanian and Turonian ammonites from the Novo
Redondo area, Angola. M.K. Howarth. 1985. Pp.
73-105. 33 figs. 0 565 07006 1.

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

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

£5.60

| Volume 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

The Cretaceous echinoid Boletechinus, with notes on the
phylogeny of the Glyphocyphidae and Temnopleuridae.
D.N. Lewis. 1986. Pp. 59-90. 11 figs. 7 tables. 0 565
07011 8.

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

£5.60
No. 3

No. 4

£13.00

Volume 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

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

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

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

Volume 42
No. 1 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 5. £46.50

Volume 43
No. 1 A Global Analysis of the Ordovician—-Silurian boundary.
Edited by L.R.M. Cocks & R.B. Rickards. 1988. 394

pp.. figs. 0 565 07020 7. £70.00

Volume 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

Cenomanian brachiopods from the Lower Chalk of

No. 2
|

Britain and northern Europe. E.F. Owen. 1988. Pp.
65-175. 0565 07022 3. £21.00

The ammonite zonal sequence and ammonite taxonomy
in the Douvilleiceras mammillatum Superzone (Lower
Albian) in Europe. H.G. Owen. 1988. Pp. 177-231. 0
565 07023 1. £10.30

Cassiopidae (Cretaceous Mesogastropoda): taxonomy
and ecology. R.J. Cleevely & N.J. Morris. 1988. Pp.
233-291. 0565 07024 X. £11.00

No. 3

No. 4

Volume 45
No. 1 Arenig trilobites—Devonian brachiopods—Triassic
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CONTENTS

105 The brachiopods of the Duncannon Group (Middle-Upper Ordovician) of southeast Ireland
M.A. Parkes

Bulletin of The Natural History Museum
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VOLUME 51 NUMBER 1 29 JUNE 1995

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]

A synopsis of neuropteroid foliage from the

Carboniferous and Lower Permian of Eurape
THE NATURA

CHRISTOPHER J. CLEAL, Brew 244434
Department of Botany, National Museum of Wales, Cardiff CF1 3NP
PRESENTED

CEDRIC H. SHUTE GENERAL LIBRARY
Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 SBD 7s

=5 JUL 1995

CONTENTS

SHIROOSIS sesseeonh dooncossesSeDUOD COMO BEDS SCP BOReOe ode OoUReCC CC MRMMRREnCoEeE Camere concn CCC RCO acer Ee eee ee cee e cette aeacr 1

RUT OAC UNO Milestone et =o Slee atest Stole Soa Soe Sse sc ee aste se = oi ic crate deseo seins Seas seeeruset oe eaten tie dase aceewscaeses 2

IMAGTINO GTS 8) oak edbedodb6 SapB eed SoCEHET PEMA eeDEC SES SHOE One e MORMeRanamEn Rt GcB 4c 8 Gr en ak AMR AN Be on aE ed Aa Ne 3

SVMOD VIM VAL StS Besser cia js Nau en clslerin wie u deat oeillasteasainsi dation tet ee reteacicteee seeded eet aeeiaee te saree eekeme ctor seamee 3
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SOUECESIOMG ata wrratees site ceivsic ins oaisiur sstis'an risiss int ose nets ore aj2 twas ee See AIRE Moree oe cele s ASS ee Tae Se ae ae ee eer 4

MAXONOMUCG DACKPLOUNG! Siicnecse sees adenine ssicws ac sesiaicioee chins viule eee eee RUMOR Renee uimaee peers ee eae ativan eaten cent eeaetees center ae 8

Gritenafor accepting alSPECieSy ..5. ssn et pact eswcee sete gs ince OR ASOT eee eo Seo nene eee ee eects 8

PNET ETETIC MOGE] Bey hie. nsmce hatte costes cole ae siete sok mcs winston a de Cra nate ame te ree pae ncteetalte Maeieme acole deena cam eee peau aeeetrdadane 8

SVSLCMAUICSM Mere e er eecinc sete st rate nt cae cencecduas cies nutseosthlncinsaneauemhucodseeuenotud some eade Ueuaen eect ebstin tenemiueetindudddesawtodaes 20

FLODUSENESS Ol SEMETICMIOGEL PE eee. ee decseke cae genie eee ax «vein deca eatineioind acc ahaneee anbiace sapeeelacee Mebane «eas eB ece REE 32

IDYNIGRSTIRY BITE NSIS, So asaa huh ecnsbatoacaé qobbc uae e Renee coc cae ABBeRCnET acc Seboduacc oc epee cscone Setiaeehbonene ce aceedeLesmecmEcREBEe Sees 32
DIVEISILVAOLMEULOPLETOIAS ASIANWHOLE; BAIA :i.leteiae Nainisle «ioealsten sR Rieelseeeieenae anehelsels saosaan cee eeteee seeesbleh ied. ceiveeneee 32
Speciesidiversityjanalysis| (Eranco-Belgian! Basim)\ee------. cee. ceeessee nee eee eee seeee ene cee eee eceee enece ae eeeneenee 33
Halaeoecolopicalicontrolsyonjspecies| distributlonsier.-1..-- see eerste eeeeereree oe eee ce eaadeniare. aes see eee eee eee eee 37
| : SPECIES GIVETSITIESHIM OLHETHATCAS/y ye. «tee hs otiesee fs de esies sgn inion siesetw lenin ns se 'eels MRA Oe acpi ets cele ostanae eine 39
Speciesidiversityjandi survivals tans. Penpryseid rate ctaemap staan qleadeteecat te secampien ab seeciasiettl acts ee ake une ansinasteaatieuidemasearc 39
AAC OPN tOSCOPLADl Vege mee ce estas ses Cea ia cue carass alts vse nijunauee te aspit ems serpeisaiosercouecdan ech tt soe actiiaceaustncecl 39

TURG GETIRIOEISS” daddscnaabeadedce aoadeobene Maan OB CUO SEE ACSAR CdS HE ae Remencadosuo antiase docu lee rnedee eG ne seCenaanraccueucuedaqunenecebome 40
IRESTIGS Bebeosduahde dec stece panera pe eES tart mic ebc cannon ca acta tElmEnaeePY nde hn daat i ndeen eh aad idcnobe bercadonercedsasoongreensu0c 41
Endemismyomind vidual form Se NCEA ep ctac ecinia fees = cestinieeenoct esas ebeseteties “vehi seine secretes sceceraces seer eee tees 41
NGuropteroidsHiromOUtSIGe BULOPE) so. 00... ceicesiacsiadseissnmnaiann genase eamenaeertins se etinerise stent setiaceniecansecenersersebee laste 41
| MBOTICIUGIN CaCI ATKS) coer eae a atge ss ctamcteaiaisis as algae en yensing eine whole Suicielee ater eeRe Reale dees neulesea- tee eae se eh lelaecietes stare ae wor eRe EEE 45
PNCKTIOW ECG CTE ICS ct sais srs ole ees to rele lees Ssisfe nelson sissies ilo sna ete secs cesta ncete eee cece eet ioeceie sae seccetesacee: 45
iSISIEINCES | SEIS abs Reee nee Coder enas UNhSooe ONee Sane SER CR doa EE BEES H BIER Aron ab ncsnhonmidan ha nBSsk -GSbE audit Sa aBnonboaasaateadgosnosaoannA 45
| Spe et es Tiniales eae iad aan ees eB eRe ae aRaee ee os Onion ona RPE eer cricicn as ohaacs cee aden oneeineescdee aacnonaoqeetianceeeadoo 51

Synopsis. The form-genus Neuropteris was initially established for compound leaves or fronds, whose pinnules had a
constricted base and a non-anastomosed venation, and which are mainly found in the Carboniferous. Using a
combination of frond/leaf architecture and cuticular features, it is now possible to divide this artificial taxonomic
concept into nine more closely circumscribed and homogenous form-genera: Neuropteris sensu stricto, Laveine-
opteris, Macroneuropteris, Margaritopteris, Neuralethopteris, Neurocallipteris, Neurodontopteris, Paripteris and
Sphenoneuropteris. In the palaeobotanical literature of the last half century (since 1940), fifty-seven adequately
circumscribed species have been identified from Europe as belonging to Neuropteris in its traditional, broad sense (a
further forty-four species names have been used, but are either based on inadequate type specimens, or have proved
to be later synonyms of other species). Of these fifty-seven ‘good’ species. fifty-one can be assigned with reasonable
| confidence to one or other of the nine form-genera mentioned above. That the classification provides a reasonably
robust expression of the natural relationships of the species is suggested by the fact that competition appears to have
been greater between species of the same form-genus than between species of different form-genera. It is possible to
correlate the distribution of some of these form-genera with the palaeoclimatic model that has been proposed based
on coal ball evidence. For instance, Neuropteris sensu stricto and Neuralethopteris appear to have belonged to plants
that favoured slightly wetter conditions within Carboniferous equatorial swamps. Laveineopteris- and Paripteris-
bearing plants seem to have been less environmentally constrained, although a change between wetter and drier
conditions seems to correlate with a change in the species present. The group as a whole seems to have been most
diverse in the peat-accumulating swamps of the Carboniferous equatorial belt, but with clear differences in the

) The Natural History Museum, 1995

species present in the paralic and intra-montane basins. In the higher southern palaeolatitudes of Gondwana, the
group is absent. In the higher northern palaeolatitudes of Angara and Kazakhstania it is also largely absent, with the

exception of some possible paripterid species.

INTRODUCTION

The study of Upper Carboniferous palaeobotany in Europe
has tended to follow two distinct lines, which may be sum-
marized as coal ball studies and adpression studies. In recent
years, the coal ball petrifactions have attracted most atten-
tion, and have yielded considerable information on the
anatomy and thereby the affinities of the plants. The import-
ance of this work is indisputable, but coal ball fossils can only
give a partial view of the Late Carboniferous equatorial
vegetation. For one thing, they only preserve plants that grew
in the peat-accumulating habitats. Although this was the
commonest habitat in the swamps, the acidic, water-logged
substrate was normally low in oxygen and nutrient, which
restricted the variety of plants it could support. Also, coal
balls only formed where sea water could percolate through
the peat deposits (Scott & Rex 1985). Where the peats
formed in a lower delta plain setting (e.g. eastern North
America, the Ukraine) there can be a good record of coal
balls, but in middle or upper delta plain settings, or intra-
montane basins, they are absent. Over much of Europe, coal
balls are restricted mainly to just one coal seam in the lower
Langsettian, with only a very few other known examples in
the Yeadonian, upper Langsettian and topmost Duckmantian
(this is excluding the silicified limnic peats in the Stephanian
and basal Permian of southern France). For a general review
of coal ball distribution, see Phillips (1980).

Of much wider occurrence in the European Upper Carbon-
iferous are plant adpressions. These preserve quite a different
part of the equatorial vegetation — mainly that growing on the
raised levee banks within the swamps. Although forming a
much smaller proportion of the original biomass, taphonomic
bias has caused them to dominate the adpression record
(Gastaldo et al. 1989). Also, because the edaphic conditions
were not as extreme, the levees supported a much more
diverse vegetation than the peat-accumulating habitats.

The abundance and diversity of the adpression assemblages
gives them considerable potential significance for under-
standing the Late Carboniferous tropical vegetation, but
there are a number of widely-perceived drawbacks. The
majority of identifiable adpressions are fragments of foliage
with (except in some ferns) little direct evidence of reproduct-
ive structures. Most ‘angiospermocentric’ neobotanists give
such foliar organs a low taxonomic status, and this attitude
has tended to rub off on palaeobotanists (‘One good fertile
specimen of a given species will tell far more than any
quantity of sterile ones’ — Andrews 1961). This viewpoint is
given support by the traditional generic taxonomy developed
by Brongniart (1822) for leaf fossils, and which is still being
used in some quarters. It is based on pinnule morphology and
venation, and is quite clearly artificial, often hiding natural
relationships and differences between species. It ignores the
fact that many of these Carboniferous leaves were architec-
turally complex structures, with many characters of potential
phylogenetic value. By viewing them holistically and incorpor-
ating such details as leaf architecture into their taxonomy, a

C.J. CLEAL AND C.H. SHUTE}

far more robust and natural classification can be developed ),
(e.g. Gothan 1941, Laveine 1967, Zodrow & Cleal 1988,
Cleal & Shute 1991a).

Another perceived difficulty with studying Upper Carbon-
iferous adpressions is that they show little anatomical detail. |
Up to a point, this is a valid criticism, at least when compared
with the quality of information that can be determined from
coal ball petrifactions. However, it should be remembered
that in many other parts of the geological column petrifac-
tions are absent or rare. It has nevertheless been possible to
determine many anatomical details from adpressions, particu-
larly of the epidermis through cuticle studies (e.g. Thomas &
Masarati 1982, Kerp 1991). Because of taphonomic factors,
such as post-mortal tectonic deformation, cuticles are not as
easy to prepare from Carboniferous adpressions as they often.
are from Mesozoic material. Nevertheless, they can some-
times be obtained from Carboniferous foliage fossils, provid- |
ing data that can be of considerable taxonomic importance
(e.g. Barthel 1961, 1962, Cleal & Zodrow 1989, Cleal &!
Shute 1991a). |

The present paper brings together the results of the
authors’ studies on one particular group of adpressions which
are particularly abundant in the Upper Carboniferous of
Europe: fragments of pteridospermous fronds that were
traditionally assigned to the form-genus Neuropteris Brongiil |
iart, and now referred to as neuropteroid fronds. By combin-'}
ing evidence of frond architecture (e.g. Gothan 1941,
Laveine 1967, Zodrow & Cleal 1988, Cleal & Shute 1991a
and epidermal structure (Barthel 1961, 1962, 1976, Cleal &
Zodrow 1989, Cleal & Shute 1991a, 1992), a revised generic
classification of the fossils was introduced by Cleal et al.
(1990). The first goal of the present study was to test the!
robustness of this classification. This was done by checking)
every species that has been recorded from Europe in the last.
half century, to see what proportion can be assigned to the)
more natural form-genera in the Cleal et al. classification. |

As a by-product of doing this check, we have built up a)
database of the geographical and temporal distribution of,
species in each of the genera. This has allowed us to see if any|
patterns can be elucidated, which may have palaeogeographi-
cal or palaeoclimatic significance. Such distributional work is, )
not novel in the Carboniferous (see Cleal 1991 for a review).|
However, by looking at the species distributions in the
context of more natural form-genera, it is believed that mo
meaningful patterns will be revealed.

These fronds mostly belong to the order of plants known asi
the Trigonocarpales (sometimes also referred to as the),
Medullosales). The order, which is only known from the).
lowland, palaeoequatorial deposits of the Carboniferous and) |.
Lower Permian, consisted mainly of shrubs and small trees.
although one small liana-like species has recently been)
described by Hamer & Rothwell (1988). They characterist- |.
ically had large dissected leaves or fronds, sometimes up to 7] P
metres long (Laveine 1986), but more typically 1-2 metreg) :
long (e.g. Cleal & Shute 1991a). In addition to the form) .
genera covered by the present study, other trigonocarpalean)
fronds include Odontopteris (Brongniart) Sternberg, 1825.

ney
a

;
|
|
i

1 metre

|
fig. 1 Reconstruction of Neuropteris heterophylla, drawn by Mrs.
Pauline Dean (prepared for part of the Evolution of Wales

Gallery, National Museum of Wales, Cardiff, See Thomas &
Cleal 1993:19).

ton Bernhardi, 1800 (see Cleal & Shute 1991b), Callipte-
‘idium (Weiss) Zeiller, 1888a, Alethopteris Sternberg, 1825,
‘Lonchopteris Brongniart, 1828, Linopteris Presl, 1838, and
eticulopteris Gothan, 1941. Traditionally, it has been
‘assumed that both the ovuliferous and microsporangiate
ctifications were attached directly to the vegetative fronds,
and a number of examples showing this have been described
‘in the literature (e.g. Dix 1932, Darrah 1937, Zodrow &
cCandlish 1980). However, there has recently come to light
2vidence that in some of the trigonocarpaleans they formed
ore or less complex strobilus-like structures, attached either

in an axillary position to the frond, or directly to the cauline
axis (Drinnan et al. 1990, Laveine et al. 1991). The individual
ovules were often large, robust structures, up to 8 cm long
(Gastaldo & Matten 1978), which probably relied on flotation
6 dispersal. The microsporangia clusters, on the other hand,
ere mainly small delicate structures, containing either

Brormow FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 3

monolete or (in the Potonieaceae) trilete prepollen (Millay &
Taylor 1979).

The group is not just important as a numerically significant
component of the Trigonocarpales (at least as represented in
the fossil record). Many species in the neuropteroid complex
have proved stratigraphically useful. Preeminent is Neuropt-
eris ovata, the base of whose range is the main index to the
base of the Westphalian D stage, but many others also play
an important role; in the biostratigraphical classification of
Carboniferous strata outlined by Cleal (1991), 2 zones and 6
subzones are named after neuropteroid species. It is thus
important to the biostratigrapher as well as the evolutionary
palaeobotanist to place the taxonomy of these fronds on a
firm footing.

METHODS

This analysis has been based on data extracted from palaeo-
botanical literature published over the last half century. A
starting date of 1940 was chosen as providing both a realistic
volume of literature to search, as well as an almost complete
cover of geographical areas yielding plant fossils in Europe.
In certain areas where there has been extensive work on
Carboniferous palaeobotany, only the most recent mono-
graphs have been used, although where necessary they have
been supplemented by other works which may document any
species omitted from the monographs. Full details of this can
be found below in the section ‘Sources of data’.

Every neuropteroid species that has been identified from
the Carboniferous and Permian of Europe during this half
century has been assessed. It has then been either:

1. Assigned to one or other of the frond form-genera out-
lined below in the section ‘Generic classification’ and,
where necessary, a new combination proposed; or

2. Assigned to the group of species that cannot be classified
in one or other of the frond form-genera; or

3. Assigned to an earlier published species as a synonym,
with brief reasons given, or a reference given to another
authority, for the proposal; or

4. Assigned to the list of nomen dubia species, that were
originally described on inadequate material.

The resulting taxonomic section of this paper thus includes
for each species that is accepted as valid (1) its name, (2) a
synonymy list (see further comments below), (3) reasons for
generic assignment, (4) any other comments, and (5) its
geographical and stratigraphical distribution.

Synonymy lists

The lists given are not complete and only include those
references that are significant for defining the species: the
basionym, the combination accepted in this analysis, where
the type specimen(s) are published if they are not included in
the protologue, and where there is a photographic record of
the type specimen(s) if the original reproduction was an
engraving or similar illustration. It also includes those species
that have been published since 1940, which are now thought
to be later synonyms.

In order to clarify the lists, they have been annotated using
a system comparable to that outlined by Matthews (1973).

However, it has been found useful to add to the range of signs
originally given by Matthews, and the full set as used here is
listed below.

* The protologue of the basionym.

‘The valid publication of the combination accepted
here.

T The type specimen(s) when not published in the proto-
logue, or photographic illustrations of them if the
original illustrations were poor.

? The inclusion of this reference is provisional due, for

instance, to poor illustration.
The present authors accept responsibility for including
this in the synonymy; if a species is included as a
synonym without the ‘.’, then it is based on another
authority, which is quoted at the end of the reference.

v_ The authors have seen the specimens in question.

Statistical analyses

The database built up as a result of this review has been
subject to statistical analysis, to try to determine distribu-
tional patterns. Univariate and bivariate statistics were calcu-
lated using the Arcus Pro-II package (version 2). The
statistics are straightforward and require little explanation
other than that the method of least-squares was used in the
regressions.

Cluster analyses were performed using the MVSP package,
on an IBM PC-AT computer. This package is particularly
useful, as it provides a routine (GSORTDATA) for showing
which species cause the clusters to form. Jaccard’s Coefficient
was used for the measure of similarity between assemblages,
as this gives no weight to cases where a particular species is
absent from both samples (Sokal & Sneath 1963). This was
deemed preferable to measures such as the Simple Matching
Coefficient, which takes such double-absences of a species
into account, and which might distort the results with local-
ities which have been only incompletely sampled. Clustering
was performed using the unweighted pair group strategy,
which on the whole tends to give a better resolution of the
clusters in binary data than the mathematically simpler single
linkage strategy (Sokal & Sneath 1963). A detailed discussion
of the relative merits of the various similarity measures and
clustering strategies available can be found in Sokal & Sneath
(1963) and Everitt (1980).

It is widely recommended (e.g. Sneath & Sokal 1973) that
similarity measures of this type should be investigated using
both cluster and ordination methods. To this end, the matri-
ces of Jaccard’s Coefficients were submitted to Gower’s
Principal Coordinates Analysis (PCO), again using the
MVSP package, which provides a series of two-dimensional
graphical plots. The results generally confirmed the patterns
observed using the cluster analysis, but did not have the merit
of such a concise graphical presentation. As they add nothing
to our conclusions, the results of these PCO analyses have not
therefore been included in the paper.

The cluster analyses were performed on matrices of binary
(presence/ absence) data for the various areas. Our informa-
tion was not really amenable to establishing quantified values
for the abundance of the species in the different areas. In any
case, it has recently been shown that such presence/absence
data in fact produce better results than quantified data in
establishing patterns of geographical distributions of plant
fossils, even where the quantified data can be reliably meas-
ured (Boulter et al. 1993).

C.J. CLEAL AND C.H. SHU

GEOLOGICAL BACKGROUND "I

Chronostratigraphical terminology

This paper is not intended as a biostratigraphical analysis.
The stratigraphical data is included only as a general guide
and is limited to the distribution between stages. For brevity!
the stage names have been abbreviated using a similai
scheme to that used by Harland er al. (1982). Unfortunately!
however, Harland et al. only used the European stages for the
Visean and Namurian; for the higher part of the Carboull
ous and the Permian, they switched to the Russian classifica-
tion. We are therefore proposing a list of abbreviations for
the full set of European stages, as summarized in Fig. 2. This
figure also gives an estimate of the duration of each of the
stages, based on the radiometric data summarized by Leeder
(1988), and which includes the dates derived from sanidine
crystals from tonsteins.

Sources of data |

The following provides a summary of the areas into which the
geographical distributional data have been divided, with a
statement as to the sources from where the palaeobotanical
information has been extracted. The locations of these areas
are plotted in Fig. 3 on a palaeogeographical map for the
Upper Carboniferous, using the same numbering of the area

as given below. The chronostratigraphical range of strata that
yield plant fossils in each of the areas is shown using the
abbreviations mentioned above. Some areas where strata of
an appropriate age are known to occur will not be foun

below. These include the Campine Basin of Belgium, th

Floha Basin of southern Germany, the North Sudetic Basin _

of the Czech Republic, the Resita and Svinita basins in
Romania, and the various basins in the Balkans. They have
been excluded from this analysis because the literature on th

plant fossils is inadequate and/or more than 50 years old.

I. South-West UK (Arn—Can). The British records hav
been divided between those south and north of the Wales
Brabant Barrier. Those from the south belong mainly to what —
Calver (1969) called the South-West Basin, and refers to th
South Wales, Forest of Dean and Bristol-Somerset coalfields —

—

~Pi_O_

Belgian Basin). The records of neuropteroid species is based!

(it excludes the Kent Coalfield, which is part of the | |

mainly on the illustrations in Crookall (1959), although his __

taxonomy has needed considerable modification (partly done —
by Laveine 1967). Some additions have also been nan
following the biostratigraphical analysis of the Welsh fossils
by Cleal (1978). |

2. Pennines (Asb—WeD). ‘This is taken in a wider sense than\
originally envisaged by Calver (1969), and includes both his
Pennines and Scottish basins. Records of plant fossils from
Scotland are relatively few but those that there are seem to

differ little from those of the Pennines. The main source of) _
data on the neuropteroid species in this area is Crookall)

(1959).

3. Franco-Belgian Basin (Pnd—WeD). This includes the
Nord-Pas-de-Calais Coalfield in northern France, and the
Mons-Charleroi-Namur Coalfield in Belgium (it also includes
the Kent Coalfield in Britain, but there are few illustrated
records of plant fossils from there). Neuropteroid species

-/NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN

il Subsystems Old Stages Old Sub-Stages

Lower Permian
Stephanian C
Stephanian B
Stephanian A

Westphalian D

Upper

Namurian B

| | Namurian C
|
|

| Namurian

Namurian A

Lower

Carboniferous |
| Visean Asb 5(2)

5)
Stage Duration of Stage in
20)

ut
Stephanian C StC 1

Westphalian C Bolsovian
ese aan Westphalian B Duckmantian
Car boniferous Westphalian A Langsettian Lan 2

|
Stephanian
| = i
|

Kinderscoutian

Alportian Alp 1
Arnsbergian 2
Pendleian CC aaa 2

Stephanian B StB
a

Westphalian D WeD
Bol

o
e
te)

NHN] wl] vd

Yeadonian Yea 1(?)

Marsdenian Mrd

vw
i)

4(?)

‘ig. 2 Stratigraphical schemes for the strata known to yield neuropteroid fossils. It includes the Heerlen set of stages and substages, the set
of stages currently accepted by the IUGS Subcommission on Carboniferous Stratigraphy, and a newly-revised set of abbreviations for the
stages. Also given is the estimated duration of each stage, based mainly on Leeder (1988).

iave been documented better in these coalfields than prob-
ibly anywhere else in the world. This is mainly due to the
‘nonographs by Stockmans (1933) and, perhaps more signifi-
_vantly, by Laveine (1967). Additional data have also been
vaken from Stockmans & Williére (1953, 1955), van Amerom
_|& Lambrecht (1979) and Paproth et al. (1983).

ft. S. Limburg (Lan-Bol). This lies between between the

NW Germany basin and the Kempe Basin of Belgium. There
_jlave been few studies on the palaeobotany of this basin in
_ecent years, the only ones with illustrations of neuropteroid

‘axa being by Jongmans (1953a, 1953b, 1954). In order to
_ tttempt a more comprehensive assessment of the fossils from
here, data has also been incorporated from Jongmans &
_/zothan (1915).

|. NW Germany (Arn—WeD). This area is based mainly
round the Ruhr Coalfield, but also includes the smaller
_}oalfields in the Osnabriick Highlands (Ibbenbiiren, Piesberg
_ nd Hiigel), which appear to belong to the same basin (Josten
_|tal. 1984). The most recent monograph on the neuropteroid
a from here is by Gothan (1953) and most of the records
juoted herein are based on this analysis. Additional records
jave been taken from Josten (1983, 1991) and Josten &
-aveine (1984).

. NE Germany (Kin-Bol). This is part of the paralic basin
hat has been discovered in deep boreholes in the region of
ostock, on the northern coast of what used to be the
-jerman Democratic Republic. Plant fossils from the
Yamurian have been documented by Kahlert (1979). The

first illustrated records of plant fossils from the Westphalian
were by Daber (1963a, 1967), but the stratigraphical informa-
tion provided is not sufficiently detailed for the purposes of
this study. More detailed evidence has recently been pro-
vided by Griindel (1992) and Kahlert (1992), and have been
used as the basis of the records incorporated in this study.
The claims that the upper part of this sequence extends up
into the Westphalian D or even Stephanian (e.g. Kahlert
1992) are based on doubtful evidence such as the presence of
conifer remains and is not accepted here.

7. Lublin (Asb-Bol). This represents the easternmost
extension of the belt of paralic deposits that extended across
northern Europe; the highest marine strata known here can
be correlated with what is known as the Vanderbeckei
Marine Band in Britain, and marks the boundary between the
Langsettian and Duckmantian stages. The best documenta-
tion of neuropteroid foliage from here is by Migier (1966),
but there are also useful but unillustrated summaries pro-
vided by Migier (1980) and Kotasowa & Migier in Bojkowski
& Porzycki (1983).

8. Zwickau-Oelsnitz (WeD). This was an intra-montane
basin formed in a small depression in present-day Saxony, SE
Germany (Pietzsch 1962). The neuropteroid taxa from here
have been documented by Daber (1955, 1957).

9. Saxony (Aut). This refers to the Erzgebirge (or Ore
Mountains), Déhlener, WeiBig and North Saxony Volcanic
basins, which lie between the Saale Trough and the Central
Bohemian Basin. They contain upper Stephanian and Rot-

GO? Upland Area
*%& Paralic Basin

# Intra-montane Basin

Fig. 3 Palaeogeographical map of Europe in the Late
Carboniferous, showing location of areas that have yielded
neuropteroid adpressions. Map based mainly on the Stephanian
reconstruction of Scotese (1986), with modifications adapted from
Bless et al. (1977) and Haszeldine (1984). The marine areas, both
shelf and deep ocean, are shaded. Locality numbers: 1 —
South-West United Kingdom; 2 — Pennines; 3 — Franco-Belgian
Basin; 4— S$. Limburg; 5 - NW Germany; 6 —- NE Germany; 7 —
Lublin; 8 — Zwickau-Oelsnitz; 9 — Saxony; 10 — Upper Silesia; 11
— Intra-Sudetic Basin; 12 — Saar-Lorraine; 13 — Alps; 14 — Massif
Central; 15 — Pyrenees; 16 —- NW Spain; 17 —N. Portugal; 18-S.
Portugal; 19— S. Spain; 20 — Svoge; 21 — Turkey; 22 — Donets; 23
—N. Caucasus.

liegende strata, although only the latter have yielded plant
fossils. The degree to which the basins were originally con-
nected is still not clear, but Barthel (1976) has shown that,
from a floristic standpoint, they combined to form a more or
less homogeneous unit. Barthel provides a brief account of
the geology, together with a detailed documentation of the
plant fossils.

10. Upper Silesia (Asb—WeD). This basin straddles the
Polish-Czech border. The name comes from the Upper
Silesia Coalfield in Poland (Bojkowski & Porzycki 1983),
while in the Czech Republic it is represented by the Ostrava-
Karvina Coalfield (Dopita & Havlena 1977). Up until the
early Namurian, it was part of the paralic belt that stretched
across northern Europe. Thereafter, however, marine influ-
ence ceased, and it became an intra-montane basin. The
neuropteroid taxa from the Namurian and lower Westphalian
of this basin are documented by Stopa (1957), Kotasowa
(1968) and Purkynova (1971). The upper Westphalian plant

C.J. CLEAL AND C.H. SHUTE

fossils are less well documented, although some useful data ig
provided by Kotasowa (1979).

11. Intra-Sudetic Basin (?Asb—Aut). This also straddles the —
Polish-Czech border. Traditionally, it was taken to include
the Lower Silesia, Podkrkonosi and Poorlicka pane
coalfields. However, sedimentological work summarized b
Holub et al. (1977) suggests that it was part of a larger area 0
sedimentation, also including the large Central Bohemia
‘basin’, as well as smaller areas of outcrop such as the
Boskovice and Blanice furrows (see also comments by Hav
lena 1953 and Wagner 1977). It is in this wider sense that we
use the term Intra-Sudetic Basin. Most of the neuropteroid
taxa are documented by Némeje (1949) and Havlena (1953).

12. Saar-Lorraine (Duc-WeD, Bar—Aut). This was a
intra-montane basin, lying between the Rheno-Hercyniar
and Saxo-Thuringian zones, and now straddling the Franco
German border. The deep borehole Saar-1 has proved thaj ©
deposition started in the late Visean (Weingart 1976). How!
ever, the exposed part of the sequence, and that which has
yielded virtually all known neuropteroid taxa, ranges fro
upper Duckmantian to Autunian, with a stratigraphical gap
from the top Westphalian D to upper Baruellian. The neu
ropteroid taxa from here have been documented by Clea
(1985) and Laveine (1989), with additional contributions by
de Jong (1974), Doubinger & Germer (1975a, 1975b))
Boersma (1978) and Cleal & Zodrow (1989). Also, althoug
it was published before the starting point that we have
selected for this study, the exceptionally illustrated mono)
graph by Bertrand (1930) cannot be ignored (although hi
species have not been included in the synonyms).

13. Alps (?Cho—Aut). Caught up in the complex tectonic —
deformation of the Alps are numerous patches of Carbonifer
ous strata yielding plant fossils. The heavy tectonism mean{
that the fossils are on the whole fragmentary and yield nq
cuticle. Also, the dislocation of the strata means that it 1
often difficult to place them in any sort of coherent strati
graphical continuum. Nevertheless, enough material ha
been collected over the years to allow many neuropteroi¢
species to be recognized from the Austrian (Fritz et al. 1990)
Swiss (Jongmans 1960) and French (Greber 1965) alps. Mos
material comes from the less tectonized Internal Zone (alsc
known as the Briangonnais Zone in France), although some —
material has also come from the External Zone.

France lie a series of mainly small, intra-montane basins tha).
developed during the Stephanian as a result of Variscay .
tectonic activity. The most important include St. Etienne (the —
eponymous area for the Stephanian Series), Autun (the
eponymous area for the Autunian Stage), Commentry, Brive
Blanzy, Bert, Decize and Decazeville; a more complete list if
provided by Doubinger & Vetter (1985). The definitive work
on the plant fossils of this area is Doubinger (1956), whe
reviewed and partially documented the upper Stephanian an¢
basal Permian palaeobotany of all of the major basins. The
main drawback of this work is that she persisted in using ¢
number of species described originally by Zeiller (1888a
1906), despite the fact that the types are totally inadequate) —
and Doubinger herself had no new material. As a conse
quence, some of these Zeiller species, which otherwise woulc

not have been included, have had to be referred to in thi! —
analysis. Other major monographs on the palaeobotany 0

14. Massif Central (Can—Aut). Within this upland area i

individual basins are by Vetter (1968 — Decazeville) and
Langiaux (1984 — Blanzy).

‘15. Pyrenees (Kin, Aut). Like the Alps, this area has been
‘subjected to considerable tectonic deformation. Terrestrial
‘Upper Carboniferous and Lower Permian strata occur in a
/number of small, isolated basins, and plant fossils are widely
‘distributed. However, there are few illustrated records of
‘them in recent years, the only significant exceptions being by
Delvolvé & Laveine (1985 — Kin) and Broutin & Gisbert
(1985 — Aut).

|
16. NW Spain (Mrd-Aut). This is an area of Upper Palaeo-
zoic deposits that ranges over parts of Asturias, Palencia and
‘Leon, and is sometimes referred to as the Cantabrian Zone.
\It was subjected to major disruption by Variscan tectonics,
resulting in sequences containing several angular unconformi-
‘ties, and preserved in a series of disjointed outcrops. Mainly
}through the work of Wagner and his collaborators, the
complex geology has been at least partly unravelled. Wagner
(1970) and Wagner & Winkler Prins (1985) provide valuable
summaries, and more detailed information can be found in
Truyols in Martinez Diaz (1983). As part of this work,
extensive collections of plant fossils have been made. Up to
the late Westphalian D, deposition was mainly marine, with
only intermittent fluvio-deltaic incursions, but at higher levels
non-marine strata become increasingly predominant. Conse-
quently, the Marsdenian to Bolsovian plant fossil record is
patchy, but from the Westphalian D upwards it is effectively
continuous into the Permian. General reviews of the plant
fossils are provided by Wagner (1959, 1962, 1966) and
Stockmans & Williére (1965), but none are complete. They
have therefore been supplemented by the records from
individual coalfields: Central Asturia (Jongmans 1952a, Wag-
ner 1971, Wagner & Alvarez-Vazquez, 1991), San Emiliano
(Moore et al. 1971), Cervera de Pisuerga (Wagner 1960, Cleal
1981), Tejerina (Wagner et al. 1969), Guardo (Wagner et al.
/1983), Cinera-Matallana (Wagner 1963, 1964), and Sabero
(Knight 1983). Also, an undocumented list of Stephanian C
fossils by Wagner & Laveine in Wagner & Martinez Garcia
(1982) has been included, being the only recent record from
_|strata of this age.

17. N. Portugal (WeD, StC—Aut). Most of the Upper Car-

doniferous and basal Permian in Portugal occurs in the north

of the country, near Oporto (Sousa & Wagner 1983). They

represent isolated intra-montane basins in the Central Iberian
tectonic zone, and according to Wagner (1983a) can be

related to the Carboniferous deposits in S. Spain (see below).
Of those containing Westphalian strata, only that at Ervedosa
qas yielded abundant plant fossils, including neuropteroid
tonds. The other basins rich in plant fossils (the Douro and
Bucaco basins) are Stephanian C to Autunian in age. The
dalaeobotany of these deposits is reviewed by Wagner &
Sousa (1983).

18. S. Portugal (WeD). This refers to three small outliers
hat are the only development of continental Upper Carbon-
ferous rocks in southern Portugal. They are the remains of
im elongate basin (the Santa Susana Basin) that developed
ilong the fracture-zone that separates the Ossa-Morena and
South Portuguese tectonic zones. Much of the sequence is
onglomeratic, but there are also coals with finer-grained
lastic deposits that have yielded plant fossils. The latter are
eviewed by Wagner & Sousa (1983).

'NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN d/

19. S. Spain (Lan, StC—Aut). Carboniferous and Permian
terrestrial deposits in the southern half of the country are
very patchy, being mainly restricted to small, fault-bounded
basins. Westphalian plant fossils have been recorded from
just two areas: the Villaneuva del Rio y Minas Coalfield in
Sevilla (Lan — Wagner ef al. 1983), and Penarroya-Bélmez-
Espinez (or Guadiato) Coalfield in Cordoba (Wagner 1983a,
1983b, 1990). A third area of Westphalian strata occurs in the
Sierra de San Pedro in Caceres (Wagner 1983a), but there
appear to be no records of plant fossils from here.

From higher strata, the best documented assemblages of
plant fossils occur near Guadalcanal in northern Sevilla
(Broutin 1986) and the Puertollano Coalfield in Ciudad Real
(Wagner 1985), In addition, there are records from Henare-
jos in Cuenca (Wagner ef al. 1985). Plant fossils have been
reported in a number of other outcrops of Autunian strata
(reviewed by Wagner & Martinez Garcia 1982, and Martinez
Diaz 1983), but none have yielded neuropteroid foliage.

20. Svoge (?Pnd—?Cho; Yea-Bol). This is the most impor-
tant coalfield in Bulgaria, and represents the remains of an
intra-montane basin (Tencov 1971). The most comprehensive
analysis of the Carboniferous plant fossils from here is by
Tencov (1977). Another major coalfield, known as the
Dobroudja Basin, has been discovered in eastern Bulgaria
below Mesozoic cover (Ten¢ov & Koulaksuzov 1972) but to
date the plant fossils have not been monographed. The
palaeobotany of the small upper Stephanian and Permian
basins in northwest Bulgaria (Tencov 1971, 1973) have also
not been revised taxonomically in recent years.

21. Turkey (Yea-WeD). Upper Carboniferous occurs in a
number of small outcrops near the northern coast of Turkey,
the most important being near Zonguldak, Amasra, Pelitova
and Azdavay. The stratigraphy is summarized by Kerey et al.
(1986), who also provide a well documented record of the
plant fossils. A more extensive listing of fossils is provided by
Jongmans (1955), but is unillustrated and so cannot be
judged.

22. Donets (Bri—Aut). The Donets Basin lies on the south-
ern edge of the Russian Platform, and has produced the most
important coalfield in eastern Europe. Brief accounts of the
Upper Palaeozoic geology of the area are given by Kler et al.
(1975) and Aizenverg et al. (1975). Prior to the very late
Visean, it was exclusively an area of marine-carbonate depo-
sition. From the Brigantian, however, deltaic complexes
frequently extended into the basin, and the rest of the
Carboniferous consists of alternating marine and non-marine
deposits. This has given the basin considerable potential
importance for correlating the so-called Heerlen chronostrati-
graphical classification, that was based on the non-marine
sequences of western Europe, and the standardized Russian
chronostratigraphy, based mainly on the marine sequences of
the Moscow Basin (Wagner et al. 1979). The most detailed
illustrated documentation of the plant fossils from here have
been by Novik (1952, 1954, 1968), although additional unil-
lustrated data are given by Fissunenko & Laveine (1984).

23. North Caucasus (?Kin—WeD, StB—StC). A number of
areas of Carboniferous outcrop occur on the northern slopes
of the Caucasus (Kavkaza) Mountains in Georgia. Their
geology is outlined by Pogrebnov (1975) and Kler et al.
(1975). Mainly Tournaisian marine deposits are overlain
unconformably by exclusively non-marine Upper Carbonifer-

ous deposits. They are of interest as the easternmost Carbon-
iferous plant-bearing deposits of Laurasia, although their
assemblages are regarded as having close affinities with those
of western Europe, closer in fact than with the geographically
nearer Donets. The plant fossils from here are described by
Novik (1952, 1978), Shchegolev (1979) and Anisimova
(1979).

TAXONOMIC BACKGROUND

Criteria for accepting a species

Most of the species listed in the nomen dubia section of this
paper are validly published according to the International
Code of Botanical Nomenclature, but in our view have been
described from insufficient material to demonstrate the range
of morphological variation. A knowledge of this variation is
essential if a species is to be usable for specimens other than
the types, and thus for it to be a viable taxonomic entity
(Cleal 1986).

There are no fixed rules for determining if a species has
been adequately defined; common sense has to be the main
guide. If it is based mainly on isolated pinnules and short
fragments of pinna, then ‘tens’ of specimens are almost
certainly needed to demonstrate the variation. If, on the
other hand, the specimens represent large segments of pri-
mary pinna branches, then the variation may be demon-
strable with less than ten. Rarely, if ever, is a single isolated
specimen a sufficient basis for describing a new species, no
matter how different it may seem to be from existing species.

The generic model

The generic classification used here has been developed from
taxonomic schemes proposed by Gothan (1941), Laveine
(1967) and Cleal et al. (1990). Those of Gothan and Laveine
were based on features of gross morphology, primarily of
frond architecture, while Cleal et al. also used epidermal
evidence. In this study, we have also used three other, less
well-known form-genera (Neurodontopteris, Sphenoneuro-
pteris, Margaritopteris) to accommodate a small number of
species, which would otherwise be unassignable. The main
diagnostic characters for each form-genus is summarized in
Table 1. In the following section, the systematics of each
form-genus is briefly summarized. It should be emphasized
that an attempt has been made to make these form-genera as
far as possible natural clusters of species, and are thus
form-genera in the sense of Cleal (1986) and Visscher et al.
(1986), rather than in the artificial sense given in the Interna-
tional Code of Botanical Nomenclature.

Form-genus LAVEINEOPTERIS Cleal, Shute & Zodrow
(1990: 489)

Type. L. loshii (Brongniart) Cleal, Shute & Zodrow

COMMENTS. This was established for the neuropteroid spe-
cies that have been shown to have large, orbicular cyclopterid
pinnules in the lower part of the frond (Figs 4, 5). Such
cyclopterids have often been taken to characterize all of the
imparipinnate neuropteroid species. As pointed out by Cleal
& Shute (1991a), however, cyclopterids are only known

C.J. CLEAL AND C.H. SHU

Fig. 4 Laveineopteris loshii (Brongniart) Cleal et al. Copy of von
Roehl (1868: fig. 17), showing orbicular cyclopterids attached
near the dichotomy of the primary rachis. Origin: Hibernia
Colliery, near Gelsenkirchen, the Ruhr, Germany. Here
reproduced at x 0-28 of original specimen.

attached to a very small range of species, all of which also
show a distinctive set of cuticular characters, such as the
virtual absence of intercellular flanges on the abaxial pinnule
surface, the absence of multicellular trichomes, and the weak
differentiation of the costal and intercostal fields of the
adaxial pinnule epidermis (Fig. 17C,D).

It is important to emphasise that the laveineopterid cyclo
pterids are different from the swollen pinnules present at the
base of the true neuropterid fronds. As pointed out by Clea
& Zodrow (1989), these cyclopterid pinnules have a mark
edly different epidermal structure from the ‘ordinary’ pin}
nules in the main part of the frond. Also, they were not}
originally orientated in the same plane as the rest of the
frond. Their function is still unclear, but it is unlikely to have ©
been simply photosynthetic.

The presence of cyclopterid pinnules suggests that Laveine
opteris is more closely related to the callipteridiums than the
neuropterids, since similar cyclopterids are known attached
to both Callipteridium and Margaritopteris (Laveine et al.
1977). Unfortunately, little is known of the epidermal struc-
ture of the callipteridiums to support this view.

The anatomy of the rachides is of a type usually associated

Soom FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN

——"??. TY eee

i = = 4 = 4 ~ (—) yisus] ajnuuid ¢-< 10} Suipuajxo pur
}UdIINIIp-uOU 10 (+) YIsUa] ajnuuId ¥-> 10F
SUIPUD}X9 puke JUoIINSap Ayyensn uldAplyy

se = = 4 = AL - puod JO aseq edu }dd9xa siyoes 0} pasny
Ajjenied Ayjewuou ore yey) sajnuulg

Z ‘ € ( % tfc b/c (-) aeuurdipenb 10 (¢) ayeuutdiy
*(Z) ajeuurdiq soysueig euuid Arewutig

= = = = = = 4 puoiy jo jaed 19MO] Ul
sajnuuld pliajdojoAd Wo}1uas 10 Ie|NdIQIO
4 = = = tL 4b - doRJINS [RIXEGR UO SOWOYOI] AP]NYJION|NYY
i = + aL ; i i - aoRjins jerxege uo aviideg

AD ({)uy (61d wy /AD Le yArs| ig/juy uy (wry) sn Ad0joAoryduue

JO (AD) aAd0]949 (1g)
on Aoviedéyoeig ‘(uy) sAd0WOUR PROS

4 = dk ae ae ae ~ sjjem [euljonue (—) ou Aypenq4ia
JO (+) Su01}s YIM JJIINNd jIexeqy
4 = de +4 = de - Jayjour duo Woy

(—) Ie IWS 10 (+) JUdIAIJIP ddRyINs
[BIXepe UO S][99 [PISOSIOJUT pur [e]SOD

+ - - = ~ = = (+) soorjins y10q 10
(—) 990RJ.IMs [eIxege A[UO UO e}PUIOIS

siajdojuopoinan sual ding Stajdoyjajvanan sidajdyjv20Anan S11a]d0Anauosrvy si4ajdoinan. si4ajdoauiaav T

SE ee ea ee Se ae
‘(1 9192.1 *Z66T) 21NYS 2 Jea[D Woy popuedxg -1aded sty} ul pod eIDUIS-WIOF dy] DsOUSEIP 0} pasn siajJoRIeYO Ie[NONO pue [RoIsojoydiow ssois oy], Tage.

— cS
CHINES
INS
TPL

Wel)
SIONS

SS
Cee

ora,

Za
nefeman'
GSS

Fig. 5 Laveineopteris rarinervis (Bunbury) Cleal et al. Copy of
Carpentier (1930: fig. 1), showing orbicular cyclopterids attached
near the dichotomy of the primary rachis. Here reproduced at x
0-7 life size.

with the Trigonocarpales (Oestry-Stidd 1979). Little is known
of the fructifications, other than that large ovules were
attached directly to the frond, probably at the end of ultimate
pinnae (Kidston 1904).

Form-genus MACRONEUROPTERIS Cleal, Shute & Zodrow
(1990: 488)

Type. M. macrophylla (Brongniart) Cleal, Shute & Zodrow

COMMENTS. Most neuropteroid fronds have a dichotomy of
the primary rachis producing tripinnate or occasionally quad-
ripinnate branches. In some species, however, the dichotomy
of the primary rachis produces less-divided, essentially bipin-
nate, primary rachis branches (Figs 6-7). These species also
have a number of distinctive epidermal characteristics, such
as brachyparacytic or cyclocytic stomata (Fig. 17E,F). It was
for this distinctive group of species that Cleal et al. (1991)
proposed the form-genus Macroneuropteris.

The fronds of Macroneuropteris are very similar to Neurop-
teris sensu stricto, except that they are less divided. Of
particular significance is the presence in at least one macro-
neuropterid species (M. scheuchzeri) of so-called ‘Odontopt-

C.J. CLEAL AND C.H. SHUTE | ~

eris lindleyana’- type pinnules (e.g. Crookall 1959: pl. 57, fig.

1), which can be compared with laciniate pinnules in the
lower part of true neuropterid fronds (e.g. Stockmans 1933:
pl. 11 fig. 1; pl. 12 fig. 2; Zodrow & Cleal 1988: pl. 4 fig. 3).

There is no evidence of the orbicular cyclopterid pinnules of |

Laveineopteris or Margaritopteris.

Nothing is known of the fructifications. Beeler (1983)
claimed that the rachis anatomy is of a type typical of the
Trigonocarpales. However, this was based purely on evi-
dence of association; she could find no such rachides with
macroneuropterid pinnules directly attached.

Form-genus MARGARITOPTERIS Gothan (1913: 168)
Type. M. coemansii (Andra) Gothan

COMMENTS. Most species included in this form-genus have
broadly attached and/or lobed pinnules, and prior to Goth-
an’s protologue were assigned to Odontopteris (Brongniart)

~— ial

i
i

Sternberg, 1825 or Sphenopteris (Brongniart) Sternberg, 1825 |
(see Laveine et al. 1977). However, one species, originally |
included in Neuropteris also belongs here (‘N.’ multivenosa |
Purkynova). Laveine et al. (1977) have shown that it is almost |
certainly the ancestral form of Callipteridium. Nothing 1s|

known of the fructifications or stem/rachis anatomy.

Form-genus NEURALETHOPTERIS Cremer ex Laveine
(1967: 97)

TyPE. N. schlehanii (Stur) Laveine

COMMENTS. This form-genus is used for alethopterid-like
fronds, in which the pinnules have a constricted base (Fig. 8).
Most of its component species were originally described as
neuropterids, but they in fact have little to do with that
form-genus in its currently defined sense.

The taxonomy of the form-genus has been thoroughly
discussed by Laveine (1967), and need not be repeated. Our
only disagreement with his analysis concerns the authorship

of the taxon. Laveine quotes Cremer (1893), but this is a]

thesis that was not effectively published. Wagner (1963, 1965)
suggested that the name should be resurrected, but provided
neither a diagnosis nor type. The first validly published
diagnosis is in fact in Laveine’s study, who must therefore be
taken as the author of the genus.

The architecture of the frond has been established with
reasonable certainty by Laveine et al. (1992). As with most of
the trigonocarpaleans, the frond had a dichotomy of the
primary rachis producing two tripinnate primary rachis
branches. Most significantly, there appear to be no interca-
lated elements on the primary rachis branches between the
secondary pinnae. The lack of this feature separates Neur-
alethopteris from most of the other neuropteroid fronds and

helps confirm that its affinities lies closest with the aletho- |

pterids.

There have been a number of reports of sporangial organs
attached or closely associated with neuralethopterid fronds
(Dix 1932, 1933; Arnold 1949; Jongmans 1954; Stockmans &
Williére 1961; Laveine 1967). Dix and Arnold both referred
them to the form-genus Aulacotheca, but Jongmans identified
them as Whittleseya. By studying a range of specimens from a
single locality, Stockmans & Williére concluded that this
apparent taxonomic difference in fact reflected infraspecific

' NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 11
}

‘ig. 6 = Macroneuropteris macrophylla (Brongniart) Cleal et al. Specimen showing lower part of frond. V.2970. Westphalian D, Radstock,

Somerset, UK. Natural size.

</
WS

OY Y Wr My px?

LU SRN

YY, LySSynWVne. Ans WY) WN
Uf, Ts WV OK \\
WY gE \} 2X\\ A

t
= FANON Mn. aN VV
ET
Ze NNN is =
Vi UNE N i.
TR ae a PO
— \ee
St
eS
sant VAN NUNN Wie Ou SDS i
NOOSE ARAN
SS, a,

NNN a
ARIS 2 u We
\\\ x wy) S 17 My
WW WW, DY SY), Wy
Ree Sy
—f
=e

Fig. 7 Reconstruction of Macroneuropteris frond (M. macrophylla
(Brongniart) Cleal et al.). Based on work done with Professor
J.-P. Laveine.

morphological variation. They therefore proposed the form-
genus Givesia for the neuralethopterid sporangial organs.

No ovules have been found attached to Neuralethopteris
fronds, although Jongmans (1954) reported large, Rhabdo-
carpus ovules in close association. There is no available
evidence of the stem or rachis anatomy.

Form-genus NEUROCALLIPTERIS Sterzel (1895: 283)

Type. N. gleichenioides (Stur) Sterzel (Neuropteris gleiche-
nioides Stur).

COMMENTS. The systematic basis of this form-genus is given
by Cleal et al. (1990). In essence, it refers to a group of
Stephanian and Lower Permian neuropteroid fronds (Fig. 9),
which have been shown to have more complex stomatal
apparatuses than typical neuropterids (Fig. 10). Relatively
little is known of the frond architecture but what information
is available (e.g. Setlik 1980) suggests that it is very similar to
that of Neuropteris sensu stricto. However, as none of the
species are that well known, the genus may not be fully
homogeneous. Evidence of fructifications or stem/rachis
anatomy is unknown.

In the upper Barruelian to Autunian is found a species
which looks very like Neurocallipteris, but has an anasto-
mosed venation. This was generally referred to as Reticulo-
pteris germarii (Giebel) Gothan, but it is now assigned to a
different form-genus, namely Barthelopteris Zodrow & Cleal
(1993). Thus, just as Reticulopteris is the mesh-veined form of
Neuropteris, and Linopteris is the mesh-veined form of Pari-
pteris, Barthelopteris is the mesh-veined counterpart of Neu-
rocallipteris.

C.J. CLEAL AND C.H. SHUTE
Form-genus NEURODONTOPTERIS Potonié (1893: 124)

Type. N. auriculata (Brongniart) Potonié

]
COMMENTS. This form-genus was originally established for |
species showing pinnule characteristics intermediate between
Neuropteris and Odontopteris. In this sense, it is clearly an |
artificial concept. However, there has been a recent recon- |
struction of the frond (Langiaux 1984: 105) from which a_
more ‘natural’ concept for the form-genus can be developed. |
Obvious characteristics include the smaller and less-divided |
frond compared with Neuropteris (Fig. 12) and the tendency i
of the pinnules to be fused to the rachis along the basiscopic ;
side. Cuticular evidence also clearly characterizes the type
{

species (described by Barthel 1976, under the incorrect name |
Neuropteris cordata — Z. Simtinek, pers. comm. 1992). Dis- |
tinctive features include the pinnules being amphistomatic !
and the cyclocytic stomata without marked papillae (Fig. 11). |
The form-genus is in clear need of revision and is used here |
only to include the type species. The frond architecture
suggests affinities with the Trigonocarpales. However, there |
is no evidence of fructifications or stem/rachis anatomy to
support this view.

Form-genus NEUROPTERIS (Brongniart) Sternberg
(1825: xi)

BASIONYM. Filicites sect. Nevropteris Brongniart (1822: 233)
Type. Neuropteris heterophylla (Brongniart) Sternberg

COMMENTS. This name was originally established by Brongn-
iart for all fossil frond fragments bearing pinnules with a
constricted base and non-anastomosed venation. Subsequent
work demonstrated that several clusters of species could be | _
recognized in the traditional concept of Neuropteris (Gothan | —
1941, Laveine 1967, Cleal & Zodrow 1989) but it was not |
certain which of them included the type species (N. hetero- |
phylla) and thus was true Neuropteris. The problem was |
solved by the study of the cuticles and frond architecture of |
the type species by Cleal & Shute (1991a), and allowed the
formal re-classification of the group by Cleal et al. (1990)
(NB. the title of the Cleal & Shute 1991a paper was changed |
at the last minute and is different to that quoted in the —
bibliography at the end of Cleal et al. 1990). It is the emended |
concept of Neuropteris proposed by Cleal et al. (1990) that is |!
used in this paper.

Despite previous preconceptions, Neuropteris sensu stricto
has pinnules that are often partly fused to the rachis, and |
have a relatively weakly developed midvein (Fig. 13). Like
Laveineopteris, the main dichotomy of the primary rachis
produces tri- or rarely quadripinnate branches (Figs 14-16).
Unlike Laveineopteris, however, there are no orbicular |
cyclopterids attached to the proximal part of the frond. |
Instead, the primary rachis below the dichotomy bears
rachides with enlarged and/or laciniate pinnules attached.
Both from their orientation relative to the rest of the frond, |

)

—

and their epidermal structures, these basal pinnules would |
seem to have simply been photosynthetic structures, not
differing significantly in function from the pinnules higher in
the frond.

During the middle Westphalian, Neuropteris developed
progressively more flexuous veins, culminating in the Bolso-
vian in a fully anastomosed venation (Josten 1962, Zodrow &
Cleal 1993). This anastomosed form of neuropterid is

1
‘

|
NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN

|
|

ig. 8 Neuralethopteris schlehanii (Stur) Laveine. V.1301. Langsettian (Westphalian A), Oldbury, West Midlands, UK. A, whole specimen,
x 1. B,C, close-ups of pinnules, x 3.

|
|
|

14 CIACLEALVANDIGHE wal

Fig. 9 Neurocallipteris neuropteroides (Géppert) Cleal et al. Richter Collection, Zwickau Museum, Germany. Lower Porphyrtuff, Planitzer
Schichten (Lower Permian), Reinsdorf, Erzgebirge, Germany (type locality). A, x 1. B-D, x 1-5. Illustrations prepared from negatives
provided by Professor. M. Barthel.

|
|

|) NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN

ig. 10 Neurocallipteris neuropteroides (Géppert) Cleal et al. Slides stored in the Museum fiir Naturkunde, Berlin. Hartensdorfer Schichten
. (Lower Permian), Hedwig Shaft (Wilde Collieries), Oelsnitz, Erzgebirge, Germany. A, adaxial cuticle, Slide No. 1/89, x 200. B,
brachyparacytic stomata on abaxial cuticle, Slide No I/89, x 500. C, papillae surrounding stomata on abaxial cuticle, Slide No II/61, x 500.

D, stomata from near the edge of an abaxial cuticle, Slide No. 1/89, x 200. Illustrations prepared from negatives provided by Professor. M.
Barthel.

Fig. 11 Neurodontopteris auriculata (Brongniart) Potonié. Czech
Geological Survey, Slide No. 226/1. Lower Stephanian B, Jivka
Member, Odolov Formation, Katetina Mine, Radvanice,
Bohemia (Intra-Sudetic Basin). A, bands of stomata in intercostal
areas, X 50. B, close-up of cyclocytic stomata, x 140.
Photographs provided by Dr Z. Simunek.

A
ONS
hy gee
Q

WU HH

Fig. 12 Reconstruction of Neurodontopteris frond. Based on
Langiaux (1984: fig. 233).

assigned to the form-genus Reticulopteris Gothan. In the
lower Westphalian D, Reticulopteris declines in abundance,
then becomes extinct to be replaced by another group of
neuropterids centred on the species N. ovata Hoffmann. The
palaeoecological background to this variation in venation is
discussed in the Diversity Analysis section, later in this paper.
Distinctive characters of the pinnule cuticles are the abund-
ant trichomes, especially on the abaxial surface, the well
developed intercellular flanges on the abaxial cuticles, and
the anomocytic or brachyparacytic stomata (Fig. 17A,B).
Beeler (1983) has demonstrated that Neuropteris sensu
stricto fronds were attached to stems belonging to the form-
genus Medullosa, providing strong support for their trigono-
carpalean affinities. Evidence as to the fructifications is less

C.J. CLEAL AND C.H. SHUTE

conclusive. Kidston & Jongmans (1911) have reported a
sporangial organ attached to a fragment of Neuropteris frond,
while Darrah (1937) and Zodrow & McCandlish (1980) have
reported ovules in apparent attachment. However, no ana-
tomical information has been obtained from these fructifica-
tions. Perhaps the most interesting point is that the ovules
seem to be attached laterally to a pinna, replacing a lateral
pinnule, whereas the laveineopterid ovules seem to have
been attached to the distal end of the pinna, replacing an
apical pinnule.

Form-genus PARIPTERIS Gothan (1941: 427)
Type. P. gigantea (Sternberg) Gothan

COMMENTS. The systematic basis of this form-genus has been
thoroughly analysed by Laveine (1967). Its distinctive
paripinnate frond architecture (paired apical pinnules, inter-
calated pinnules on the penultimate rachides) separates it
from all of the other neuropteroid form-genera (Fig. 16).

There is a mesh-veined counterpart of Paripteris, known as
Linopteris Presl. It would seem that it is the foliage of a
distinctive group of trigonocarpalean pteridosperms, which
may be referred to as the Potonieaceae (see Cleal, 1993). In
addition to the distinctive frond architecture, at least one
member of the family (Linopteris obliqua Bunbury) has been
shown to have stems with a vascular system that is not as
dissected as in the other trigonocarpaleans, and when pre-
served as a petrifaction is known as Sutcliffia (Stidd et al.
1975). The ovules are generally assumed to be of the type
known as Hexagonocarpus (or Hexapterospermum when pre-
served anatomically), and are characterized by a six-fold axial
symmetry (Taylor 1966), in contrast to the three-fold sym-
metry of other trigonocarpalean ovules. Perhaps most distine-
tive are the male fructifications, which consist of numerous
sporangial clusters (individually known as Potoniea) formed
into a large cone-like structure (Laveine ef al. 1991). They
contain trilete prepollen, in contrast to the monolete prepol-
len of the other trigonocarpaleans (Stidd 1978).

The morphological evidence for the distinctiveness of the
Potonieaceae is also supported by its distribution (Laveine et
al. 1989). The Potonieaceae originated in the Visean of China
and did not appear in Laurasia until the Namurian. The rest
of the Trigonocarpales, in contrast, seem to have originated
in Laurasia and only a few species are found in China. Most
authors still retain the Potonieaceae in the Trigonocarpales,
but there is increasing evidence that it represents a totally
distinct group of pteridosperms, the few similarities (e.g.
detailed ovule structure) being a matter of analogy.

Form-genus SPHENONEUROPTERIS Shchegolev
(1979: 158)

Type. S. elegans Shchegolev

COMMENTS. This refers to a group of mainly Stephanian
fronds that stand apart from most other neuropteroids, in
having large, relatively lax pinnules with a low vein density.
Wagner (1963) and Knight (1983) put forward evidence to
show that at least some of the species (‘N.’ dimorpha, ‘N.|
praedentata, ‘Mixoneura’ wagneri) cluster together to form a) «
more natural group, although they did not propose a new
name for the group.

ig. 13 Neuropteris obliqua (Brongniart) Zeiller. V.63723. Duckmantian (Westphalian B), Rhigos, near Hirwaun, Mid-Gla
| whole specimen, xX 1. B, enlargement of pinnules, x 2.

Paripteris pseudogigantea (Potonié) Gothan. V.63724. Duckmantian (Westphalian B), Rhigos, near Hirwaun, Mid-Glamorgan, UK. C,
| whole specimen, x 1. D, enlargement of pinnules, x 3.

18 C.J. CLEAL AND C.H. SHUTE

Fig. 14 Neuropteris obliqua (Brongniart) Zeiller. Duckmantian (Westphalian B), Yorkshire, UK. Photograph taken in the field of the
proximal portion of a frond preserved in sandstone. Previously illustrated at lower magnification by Scott (1978: pl. 27, fig. 1). A, whole
specimen, X 0-2. B, pinnate foliage from above dichotomy, x 0-5. C, pinnae attached to primary rachis below the dichotomy, x 1.

'NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN

‘ig. 15 Neuropteris heterophylla (Brongniart) Sternberg. V.1797. Duckmantian (Westphalian B), Clay Cross, Derbyshire, UK. x 0-34.

Pr)
aoa ot,
Us on, f

24,

Fig. 16 Reconstruction of Neuropteris frond (N. heterophylla
(Brongniart) Sternberg). From Cleal & Shute (1991: fig. 29).

In his investigations on Stephanian plant fossils from the
Caucasus, Shchegolev (1979) described some fragments of
neuropteroid fronds, which also had relatively large, lax-
limbed pinnules and wide venation, and for which he pro-
posed the new name Sphenoneuropteris. We still have very
little information on the architecture of these fronds, and
nothing of the epidermal structure or fructifications. It is far
from clear, therefore, whether this is a homogeneous group
of species. However, for the time being Sphenoneuropteris
provides a convenient receptacle for these distinctive frond
fragments, which clearly have little to do with Neuropteris
sensu stricto, or probably even the Trigonocarpales in gen-
eral.

SYSTEMATICS

Form-genus LAVEINEOPTERIS Cleal, Shute & Zodrow

Laveineopteris guadiatensis (Wagner) Cleal & Shute,
comb nov.

*1983b Neuropteris guadiatensis Wagner: 95; pl. 1.

REASON FOR GENERIC ASSIGNMENT. Wagner records associ-
ated orbicular cyclopterid pinnules with the more typical
pinnate foliage of this species. Also, fragmentary cuticles
prepared by C.R. Hill (Natural History Museum) and shown
to us, display a number of laveineopterid characteristics:
adaxial cuticle shows relatively uniform cell patterns, no
anticlinal walls preserved on abaxial cuticle, and no trichomes
are preserved on either cuticle.

OCCURRENCE. S. Spain (Duc).

C.J. CLEAL AND C.H. SHUTE

Laveineopteris hollandica (Stockmans) Cleal & Shute, |
comb. nov.

* 1933 Neuropteris hollandica Stockmans: 31-34; pl. 10,
fig. 1. .

v 1959 Neuropteris rytoniana Kidston ex Crookall:
113-114; pl. 52, figs 3-4; pl. 54, fig. 1 (vide Laveine,
1967).

.v 1959 Neuropteris formosa Kidston ex Crookall: 139-140;
pl. 52, figs 1-2.

REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule
morphology with L. tenuifolia.

COMMENTS. Although its venation is not entirely typical, N.|_
formosa is taken to be a later synonym of L. hollandica based
on the similarity in shape of its subtriangular pinnules. Also,
the types of N. formosa originated from the same locality as|_
the types of N. rytoniana, which Laveine (1967) assigned to
L. hollandica.

OCCURRENCE. Pennines (Lan-Bol), Franco-Belgian Basin}
(Lan-Duc), S. Limburg (Lan), NW Germany (Lan-Bol), NE |
Germany (Lan-Bol).

Laveineopteris jongmansii (Crookall) Cleal & Shute,
comb. nov.

v? 1888 Neuropteris plicata Sternberg; Kidston: 313; pl. 1,
fig. 1.

? 1917 Neuropteris subplicata Kidston: 1031.

*v 1959 Neuropteris jongmansii Crookall: 178; pl. 51, fig. 1.

.v 1967 Neuropteris chalardi Laveine: 176-181; pls 35-39.

REASON FOR GENERIC ASSIGNMENT. Great similarity of pin-
nule morphology to L. tenuifolia.

COMMENTS. Laveine (1967) noted the close similarity
between his N. chalardi and the holotype of L. jongmansii
figured by Crookall (1959). Crookall’s specimen alone was
inadequate evidence for Laveine to make a proper compari- |
son. However, one of us (CJC) has examined additional
material in the collections of the British Geological Survey
and can confirm that the two species are identical.

The type and only known specimen of N. subplicata has
similar shaped pinnules and a dense venation. The apical |
pinnule is rather small, but can be compared with the lower |
end of the range of variation of L. jongmansii (e.g. Laveine |
1967: pl. 37, fig. 2). More examples of this species are needed |
but, if the synonymy can be confirmed, Kidston’s species will |
be the valid name.

OCCURRENCE. Pennines (Bol), Franco-Belgian Basin (Bol),
NW Germany (Bol), Lublin (Bol), NE Germany (Bol).

Laveineopteris loshii (Brongniart) Cleal, Shute &
Figs 4, 17C,D

* 1831 Nevropteris Loshi Brongniart: 242; pl. 72, fig. 1; pl.
73.

.v 1959 Neuropteris hemingwayi Crookall: 121-122; pl. 46,
fig. 6.

T 1967 Neuropteris loshi Brongniart, Laveine: pls C-D.

S 1990 Laveineopteris loshii (Brongniart) Cleal, Shute &
Zodrow: 490.

Zodrow

REASON FOR GENERIC ASSIGNMENT. Type species.

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 7)

ig. 17 Neuropteroid cuticles photographed using Normarski Interference. All x 200. A, Neuropteris ovata Hoffmann. Abaxial cuticle.
V.62925. Basal Westphalian D, Kallenberg Seam, Itzenplitz Colliery, Saarland, Germany. B, N. ovata. Adaxial cuticle. V.62924. Same
horizon and locality. C, Laveineopteris loshii (Brongniart) Cleal et al.. Abaxial cuticle. V.62974. Duckmantian (Westphalian B), Royosborn
Colliery Borehole, North Yorkshire, UK. D, L. loshii. Adaxial cuticle. V.62948. Same horizon and locality. E, Macroneuropteris
macrophylla (Brongniart) Cleal et al.. Abaxial cuticle. V.62295. Upper Westphalian D, Upper Bonnar Seam, Brogan’s Pit, Sydney
Coalfield, Cape Breton, Canada. F, M. macrophylla. Adaxial cuticle. V.63055. Same horizon and locality.

COMMENTS. The epidermal structure has been described by
Cleal & Shute (1992) and orbicular cyclopterid pinnules have
been shown attached near the base of its frond (von Roehl
1868: pl.17).

N. hemingwayi was based on a single fragment from the
Parkgate Coal (upper Langsettian) of Yorkshire, from which
L. loshii is well documented (e.g. Crookall 1959: p1.28, fig.4).
It is poorly preserved, but the pinnule shape and venation
seem indistinguishable from L. loshit.

OCCURRENCE. South-West .UK (Lan-Bol), Pennines, UK
(Lan-Bol), Franco-Belgian Basin (Lan-Bol), S. Limburg
(Lan), NW Germany (Lan-Bol), NE Germany (Lan-Bol),
Lublin (Lan-Bol), Intra-Sudetic Basin (Duc-Bol), U. Silesia
(Lan-Bol), Svoge (Duc-Bol), Donets (Lan-Duc).

Laveineopteris morinii (Bertrand ex Laveine) Cleal &
Shute, comb. nov.

* 1967 Neuropteris morinii Bertrand ex Laveine: 227-228;
pls 63-65.

REASON FOR GENERIC ASSIGNMENT. The association of
orbicular cyclopterid pinnules (Laveine 1967: pl. 64 fig. 6),
and general similarity of some of the pinnules to the more
elongate-pinnule laveineopterids such as L. tenuifolia and L.
hollandica.

COMMENTS. Laveine (1967) argued that this species belongs
to the general group allied to Neuropteris obliqua, and would
thus be retained in Neuropteris in its restricted sense as used
here. This was based mainly on the supposed presence of
forma impar-type pinnules. However, the best example that
he illustrates to justify this opinion (Ibid. pl. 65 fig. 5) is
poorly localized and there is no evidence that it was associ-
ated with more typical pinnules of this species. The other two
examples (Ibid. pl. 63 figs 2-3), although in clear association
with specimens showing the more typical pinnule form of this
species, are isolated pinnules — one possibly a terminal, the
other a lateral. Being isolated, it is far from certain that they
are of the forma impar type from the lower part of a frond, or
even that they belong to the same species.

It is true that the cyclopterid illustrated by Laveine is also
only associated with the specimens of pinnate foliage. How-
ever, in view of the close similarity of the pinnules to L.
tenuifolia (from which it can only be reliably distinguished by
its denser, occasionally flexuous veins), we believe that the
association with the cyclopterid reflects an original organic
connection.

OCCURRENCE. Franco-Belgian Basin (Bol).

Laveineopteris nicolausiana (Gothan) Cleal & Shute,
comb. nov.

* 1913 Neuropteris nicolausiana Gothan: 213; pl. 48; pl. 49,
fig. 1.

REASON FOR GENERIC ASSIGNMENT. The similarity of the
pinnule shape to L. rarinervis and of the venation to L.
tenuifolia. Also, the frequent association of orbicular cyclop-
terids.

COMMENTS. Many authors have regarded this as indistin-
guishable from L. rarinervis (e.g. Stockmans 1933, Crookall
1959, Laveine 1967). However, it differs from that species in
having (a) more linguaeform lateral pinnules, (b) lateral veins

C.J. CLEAL AND C.H. SHUTE

that fork at a narrower angle and meet the pinnule margin at ;
a more oblique angle, and (c) smaller, more ovoid apical’
pinnules. It is thus in some ways morphologically intermedi-
ate between typical L. rarinervis, and the larger-pinnuled
species L. tenuifolia. In view of its stratigraphical occurteiag
at the lower end or just below the range of L. rarinervis, it is)
possible that it represents its evolutionary ancestor and a |
with the larger-pinnuled L. tenuifolia group of species.
Bertand (1930) altered the spelling to nikolausii, changing
it to the substantive form and reverting to the original spelling
of the surname Nikolaus. However, ICBN Article 73 allows a
species name to be based on an latinized personal name in am
.

adjectival form. The original spelling is therefore retained.

OCCURRENCE. Intra-Sudetic Basin (Bol), U. Silesia (Duc-
Bol), Saar-Lorraine (Duc-Bol).

Laveineopteris piesbergensis (Gothan) Cleal & Shute,
comb. nov.

* 1953 Imparipteris piesbergensis Gothan: 57; text fig. 8; pl.
a2)

REASON FOR GENERIC ASSIGNMENT. The presence of orbicu-
lar cyclopterids in the proximal part of the frond (Gothan
1953: text fig. 8), and the close similarity of the pinnules to L.
rarinervis.

COMMENTS. Gothan distinguished this species from L. rari-
nervis by a number of characters of the orbicular cyclopteri |
pinnules, which are probably of doubtful taxonomic signifi-
cance. However, he also mentioned that the venation of the
lateral pinnules was denser and more oblique to the ies )
margin. The significance of these differences of veinin,
pattern is not clear, and Laveine (1967) included Gothan’s
species in the synonymy of L. rarinervis. However, we hae
opted to maintain the distinction, at least until the Germa
material can be more fully assessed.

OCCURRENCE. NW Germany (WeD).

Laveineopteris rarinervis (Bunbury) Cleal, Shute & |
Zodrow Fig.

* 1847 Neuropteris rarinervis Bunbury: 425; pl.22.
51990 Laveineopteris rarinervis (Bunbury) Cleal, Shute §

Zodrow: 490. .

REASON FOR GENERIC ASSIGNMENT. Epidermal structure
(Cleal & Zodrow 1989) and the presence of orbicular cyclop; ~
terid pinnules near the dichotomy of a bipartite .
(Laveine 1967: pl.45, fig.3).

OCCURRENCE. South-West UK (Bol-Can), Pennines (Bol
WeD), Franco-Belgian Basin (Bol-WeD), NW Germany |
(Bol-WeD), NE Germany (Bol), Lublin (Duc-Bol), U. Sile
sia (Bol-WeD), Donets (Duc-WeD), Turkey (WeD), (?)N
Spain (WeD).

}

Laveineopteris tenuifolia (Sternberg) Cleal, Shute &
Zodrow

T 1820 Filicites tenuifolius Schlotheim: 405; pl. 22, fig. 1.

* 1825 Neuropteris tenuifolia Schlotheim ex Sternberg: xvil

* 1990 Laveineopteris tenuifolia (Sternberg) Cleal, Shute
Zodrow: 490.

‘WNEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 23

‘REASON FOR GENERIC ASSIGNMENT. Epidermal structure
‘(Barthel 1962, Cleal 1985). Also, pinnate fragments of this
\species are almost invariably associated with orbicular cyclop-
terid pinnules.

(COMMENTS. This species has been widely reported from the

‘Iberian Peninsula. However, Cleal (1981) analysed these
records and showed that they were based either on specimens
lof Neuropteris resobae Cleal (q.v.), or on unidentifiable
‘fragments; L. tenuifolia would seem to be absent from this

area.

‘OCCURRENCE. South-West UK (Lan-WeD), Pennines (Lan-
Bol), Franco-Belgian Basin (Lan-WeD), S. Limburg (Lan),
NW Germany (Lan-WeD), Lublin (Lan-Bol), Intra-Sudetic
Basin (Duc-Bol), U. Silesia (Lan-Bol), Saar-Lorraine (Bol),
Svoge (Duc-Bol), Donets (Duc-WeD).

Form-genus MACRONEUROPTERIS Cleal, Shute & Zodrow
Macroneuropteris britannica (Gutbier) Cleal, Shute &

_ Zodrow

* 1835 Odontopteris britannica Gutbier: 68, pl. 9, figs 8-11.
* 1990 Macroneuropteris britannica (Gutbier) Cleal, Shute
& Zodrow: 488.

REASON FOR GENERIC ASSIGNMENT. Epidermal structure
(Barthel 1962).

OCCURRENCE. Zwickau-Oelsnitz (WeD).

Vacroneuropteris macrophylla (Brongniart) Cleal,
Shute & Zodrow Figs 6-7 17E-F

1831 Nevropteris macrophylla Brongniart: 235; pl. 65, fig.
| .

* 1990 Macroneuropteris macrophylla (Brongniart) Cleal,
. Shute & Zodrow: 488.

REASON FOR GENERIC ASSIGNMENT. Type species.

COMMENTS. Cuticles have been described by Cleal & Zod-
“ow (1989). The frond architecture is currently under review
py Cleal, Laveine & Shute.

| The specimens from North Caucasus illustrated by Anisi-
nova (1979) as this species are clearly misidentified. They are
solated pinnules which resemble those of Paripteris (e.g. P.
seudogigantea), although they would seem to have origi-
nated from rather a high stratigraphical position (WeD) for
hat form-genus.

ICCURRENCE. South-West UK (WeD-Can)

Macroneuropteris scheuchzeri (Hoffmann) Cleal, Shute
& Zodrow

1827 Neuropteris scheuchzeri Hoffmann: 157; pl. 1b, figs
144.

1990 Macroneuropteris scheuchzeri (Hoffmann) Cleal,
Shute & Zodrow: 488.

REASON FOR GENERIC ASSIGNMENT. Epidermal structure
‘Barthel 1961, Cleal & Zodrow 1989). The frond architecture
S currently under review by Cleal & Laveine.

i= Havlena (1953: pl. 5, fig. 3) figured a fragment

reputedly from the Stephanian C of the Intra-Sudetic Basin as
Neuropteris cordata. He claimed similar material also
occurred in the Autunian of this region. As pointed out by
Laveine (1967), however, the figured specimen is almost
certainly M. scheuchzeri. If it does belong there and its stated
provenance is correct, this is by far the highest stratigraphical
occurrence of this species in Europe.

OCCURRENCE. South-West UK (Duc-Can), Pennines (Lan-
Bol), Franco-Belgian Basin (Duc-WeD), S. Limburg (Duc),
NW Germany (Duc-WeD), NE Germany (Duc-Bol), Lublin
(Bol), Intra-Sudetic Basin (Duc-WeD, ?StC-Aut), U. Silesia
(Lan-Bol), Saar-Lorraine (Bol), Alps (Can), Svoge (Bol),
Donets (Duc-WeD), N. Caucasus (WeD), Turkey (WeD), N.
Portugal (WeD), NW Spain (WeD-Can), S. Portugal (WeD).

Macroneuropteris subauriculata (Sterzel) Cleal, Shute
& Zodrow

T 1855 Neuropteris auriculata Brongniart: Geinitz: pl. 27,
figs 4-7, 9.

1901 Neuropteris subauriculata Sterzel: 100.

1990 Macroneuropteris subauriculata (Sterzel) Cleal,
Shute & Zodrow: 488.

*
§

REASON FOR GENERIC ASSIGNMENT. Epidermal structure
(Barthel 1962) and possibly frond architecture (Daber 1957).

COMMENTS. This species appears to be endemic to the
Zwickau-Oelsnitz Basin. The record from North Caucasus by
Anisimova (1979) is based on extremely poorly preserved
material and is unconvincing.

Laveine (1989: pl. 60, fig. 1) has figured a single fragment
of this species from the upper Westphalian D (or possibly
basal Cantabrian) of Saar-Lorraine. It bears a close similarity
to the fragmentary types of Neuropteris germeri de Jong,
from slightly older strata in Saar-Lorraine, and which we
have provisionally assigned to N. ovata. Cuticles from this
Saar-Lorraine material could help resolve their taxonomic
position.

OCCURRENCE. Zwickau-Oelsnitz (WeD).

Form-genus MARGARITOPTERIS Gothan

Margaritopteris multivenosa (Purkynova) Cleal &
Shute, comb. nov.

1970 Neuropteris multivenosa Purkynova: 223-224; pl.
45, fig. 1, pl. 46, fig. 1.

REASON FOR GENERIC ASSIGNMENT. The prominent midvein
and the size and texture of the pinnules (see Laveine et al.
1977). Also the manner of lobing of the pinnules.

COMMENTS. Laveine et al. (1977) were clearly of the opinion
that this species belongs to Margaritopteris, and was the
precurssor of the more familiar Westphalian species,
although no formal proposal of transference was made. The
factors mentioned above, particularly well shown in Laveine
et al. (1977, pl. 19, fig. 3), make it unnecessary to postpone
the proposal of transference.

OCCURRENCE. U. Silesia (Alp).

24
Form-genus NEURALETHOPTERIS Cremer ex Laveine

Neuralethopteris densifolia Josten

* 1983 Neuralethopteris densifolia Josten: 144; pl. 53, fig. 1;
pl. 54, fig. 1.

REASON FOR GENERIC ASSIGNMENT. The alethopteroid
nature of the venation.

OCCURRENCE. NW Germany (Kin- Yea).

Neuralethopteris doubravica (Purkynova) Cleal &
Shute, comb. nov.

* 1971 Neuropteris doubravica Purkynova: 165-166; pls
6-9.

REASON FOR GENERIC ASSIGNMENT. The cordate base of the
pinnules and the prominent midvein.

COMMENTS. The generic position of this species is far from
certain. At least some of the specimens figured in the
protologue, particularly those with smaller pinnules,
approach Neuropteris in venation and pinnule shape (e.g. pl.
8, fig. 1a). As pointed out by Purkynova, however, the larger
pinnules share many characters with Neuralethopteris, espe-
cially N. jongmansii, and so we propose to transfer the
species there.

OCCURRENCE. U. Silesia (Lan).

Neuralethopteris jongmansii Laveine
* 1967 Neuralethopteris jongmansii Laveine: 107; pls 2-4.

REASON FOR GENERIC ASSIGNMENT. The typically alethop-
terid nature of venation (Laveine 1967). Also, Whittleseya
sporangial structures and Trigonocarpus ovules have been
linked by Jongmans (1954) to foliage identified as N. jong-
mansii by Laveine (1967).

OCCURRENCE. South-West UK (Lan), Pennines (Lan),
Franco-Belgian Basin (Lan), S. Limburg (Lan), NW Ger-
many (Lan), U. Silesia (Lan).

Neuralethopteris larischii (Susta) Laveine

‘ 1930 Neuropteris Larischi Susta: 5, pl.1.
’ 1967 Neuralethopteris larischi (Susta) Laveine: 102; pl. 1.

REASONS FOR GENERIC ASSIGNMENT. Typically alethopterid
nature of venation (Laveine 1967).

OCCURRENCE. Franco-Belgian Basin (Arn-Lan), NW Ger-
many (Arn-Lan), U. Silesia (Kin-Lan), N. Caucasus (?Kin-
?Yea), Turkey (Yea), NW Spain (Lan).

Neuralethopteris neuropteroides (Susta) Josten

* 1927 Alethopteris neuropteroides Susta: 4; pl. 1, fig. 2.
* 1983 Neuralethopteris neuropteroides (Susta) Josten: 138;
pl. 50, fig. 1.

REASON FOR GENERIC ASSIGNMENT. The close similarity in
pinnule shape to N. schlehanii, and the tendency of the
pinnules to be fused to the rachis at the base.

COMMENTS. This species seems to occupy a position interme-
diate between Neuralethopteris and true Alethopteris.

C.J. CLEAL AND C.H. SHUTE
OCCURRENCE. Franco-Belgian Basin (Kin-Lan), NW Ger-
many (Kin-Lan), U. Silesia (Yea-Lan).
Neuralethopteris rectinervis (Kidston) Laveine

*v 1888
T 1959

Neuropteris rectinervis Kidston: 314; pl.1, figs 2-4.
Neuropteris Schlehani forma rectinervis (Kidston)
Crookall: 145-147; pl. 35, figs 6-8.
Neuralethopteris rectinervis (Kidston) Laveine: 120;
pl. 9.

8 1967

REASON FOR GENERIC ASSIGNMENT. typically alethopterid
nature of venation (Laveine 1967).

OCCURRENCE. South-West UK (Lan), Pennines (Lan),
Franco-Belgian Basin (Lan), NW Germany (Lan), U. Silesia
(Lan), Donets (Lan).

Neuralethopteris schlehanii (Stur) Laveine

NST
1953

Neuropteris Schlehani Stur: 289; pl.28, figs 7-8.
Neuropteris schlehanioides Stockmans & Williére:
233; pl. 31, figs 3, 7; pl. 36, fig. 2.

Neuropteris loriformis Stockmans & Williére: 234;
pl. 16, fig. 2.

Neuropteris rectinervis forma obtusa Tenéov: 59-60;
pl. 20, figs 3-4.

Neuropteris lata Tenéov: 60; pl. 21, figs 2-3.
Neuropteris longifolia Tenéov: 61; pl. 21, figs 4-9.

oP IPSS)
1977

1977
1977

REASON FOR GENERIC ASSIGNMENT. Type species.

COMMENTS. Epidermal structures have been described by
Cleal & Shute (1992). Also, Aulacotheca sporangial struc-
tures were reported in close association with it by Laveine
(1967: pl.5, fig.3).

The types of N. schlehanioides clearly represent fragments |

from high in the pinna of N. schlehanii and can be compared

with parts of Laveine (1967: pl. 6, figs 2-3). The type of N. |

loriformis is less typical, having extremely long, slender
pinnules. As pointed out by Stockmans & Williére (1953),
however, it occurs in association with N. schlehanii and has a
comparable venation pattern.

Neuropteris longifolia and N. lata were erected for speci-
mens from the Svidnaya Formation of the Svoge Basin. The
pinnules are rather large (up to 30 mm long), but are
otherwise very similar to N. schlehanii, with which they are
closely associated. As Laveine (1967: pl. 8) has figured
specimens of N. schlehanii with pinnules approaching these in
size, there seems little reason for separating these species.

Tenéov (1977) described the types of N. rectinervis forma
obtusa as having a venation nearer to that of N. rectinervis
than N. schlehanii. However, the veining is in fact quite
compatible with N. schlehanii, being broadly arched (cf.
Laveine 1967: pl. 7 fig. 1; pl. 8 fig. 4).

OCCURRENCE. South-West UK (Lan), Pennines (Lan),
Franco-Belgian Basin (Pen-Lan), S. Limburg (Lan), NW
Germany (Mrd-Lan), NE Germany (Kin-Lan), (?)Lublin
(Kin-Yea), Intra-Sudetic Basin (Lan), U. Silesia (Alp-Lan),
Svoge (Yea-Lan), Donets (Pen-Lan), N. Caucasus (?Kin-
Lan), Turkey (Lan), Alps (Lan), Pyrenees (Kin), NW Spain
(Lan).

Fig. 8 |

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 25

Form-genus NEUROCALLIPTERIS Sterzel

Neurocallipteris gallica (Zeiller) Cleal & Shute, comb.
nov.

* 1888a Neuropteris gallica Zeiller: 248; pl. 29, figs 1-3.

REASON FOR GENERIC ASSIGNMENT. Close similarity to N.
planchardii in both pinnule shape and venation.

_ COMMENTS. This species is not well documented, and it is not
entirely certain that it is distinct from N. planchardii. As

pointed out by Wagner (1963), many of the differences

_ claimed by Zeiller may be merely a function of the position of

_the pinnules within a frond. The most obvious difference is

_ the presence of hairs near the midvein of N. gallica, but this
could be influenced by taphonomic factors. There have,

however, been a number of records of the species in recent
years, and so it has provisionally taken to be ‘good’.

OCCURRENCE. Saar-Lorraine (Bar), Massif Central (StC),
NW Spain (StB), S. Spain (StC), N. Portugal (StC).

' Neurocallipteris neuropteroides (G6ppert) Cleal, Shute
& Zodrow Figs 9-10
:

-* 1836 Gleichenites neuropteroides Goppert: 186; pls 4-5.
* 1990 Neurocallipteris neuropteroides (Géppert) Cleal,
Shute & Zodrow: 489.

| REASON FOR GENERIC ASSIGNMENT. Type species.

COMMENTS. Epidermal structure (Barthel 1962, 1976) and
frond architecture (Barthel 1976, Setlik 1980) have been
| documented for this species.

OCCURRENCE. Saxony (Aut), Intra-Sudetic Basin (StB-
| Aut), Massif Central (StC), Pyrenees (StC), NW Spain (StC),
'N. Portugal (StC-Aut).

Neurocallipteris planchardii (Zeiller) Cleal, Shute &
Zodrow

'* 1888a Neuropteris planchardii Zeiller: 246; pl. 28, figs 8-9.
* 1990 Neurocallipteris planchardii (Zeiller) Cleal, Shute &
Zodrow: 489.

IREASON FOR GENERIC ASSIGNMENT. Similarity of epidermal
structures to N. neuropteroides (see Reichel & Barthel 1964,
Barthel 1976)

OCCURRENCE. Saxony (Aut), Intra-Sudetic Basin (Aut), N.
Caucasus (StC), Alps (?Can), Massif Central (StB-Aut),
Pyrenees (StC), NW Spain (WeD-StB), N. Portugal (StC-
Aut).

Form-genus NEURODONTOPTERIS Potonié

|
}

Veurodontopteris auriculata (Brongniart) Potonié Figs
11-12

? 1830 Nevropteris auriculata Brongniart: pl. 36.

1831 Nevropteris auriculata Brongniart: 236.

1831 Nevropteris dufrenoyi Brongniart: 246.

_ 1893 Neurodontopteris auriculata (Brongniart) Potonié:
124.

.(?)1937 Neuropteris densinervosa (Grigoriev) Zalessky: 183;
fig. 31.

REASON FOR GENERIC ASSIGNMENT. Type species.

COMMENTS. There have been no descriptions of cuticles
under this species name. However, Z. Simunek (pers. comm.
1992) has shown that cuticles described by Barthel (1976) as
Neuropteris cordata in fact belong to Neurodontopteris
auriculata. In contrast to typical N. cordata from the Massif
Central, the specimens which yielded the cuticles have
smaller pinnules (<60 mm long) with a rounder apex, and a
denser venation (30 veins per cm on the pinnule margin).
Siminek has prepared very similar cuticles from specimens of
N. auriculata from the Intra-Sudetic Basin, examples of
which are shown in Fig. 11.

The inclusion of N. densinervosa here must be regarded as
tentative, since only two specimens have been illustrated in
the literature and one of those (the holotype) only as a
drawing. However, the large pinnules and high stratigraph-
ical position (Gzhelian) of the specimens would seem com-
patible with N. auriculata.

OCCURRENCE. Saar-Lorraine (Aut), Saxony (Aut), Intra-
Sudetic Basin (StB-Aut), NW Spain (StB), S. Spain (StC-
Aut).

Form-genus NEUROPTERIS (Brongniart) Sternberg

Neuropteris antecedens Stur

* 1875 Neuropteris antecedens Stur: 53; pl.15, figs 1-6.
1953 Neuropteris mathieui Stockmans & Williére: 227.
1955 Neuropteris condrusiana Stockmans & Williére: 12;
pl. 6, figs 1-7, 9-15; pl. 9, figs 1-8.

1955 Neuropteris papilioniformis Stockmans & Williére:
13; pl. 2, fig. 3.

1955 Neuropteris pseudozamites Stockmans & Williére:
13, pl. 2, fig. 1.

REASON FOR GENERIC ASSIGNMENT. The tendency of the
pinnules to be broadly attached to the rachis and on the
venation sometimes being flexuous (Crookall 1959).

COMMENTS. The type of N. mathieui was found associated
with typical specimens of N. antecedens, and it is difficult to
see why it is not merely the small-pinnuled form of that
species.

The types of N. condrusiana, N. papilioniformis and N.
pseudozamites all originated from the same horizon and
locality, and were associated with a specimen which Stock-
mans & Williére (1955) identified as their N. mathieui. All of
this material clearly belongs to a single species, and bears
quite a striking similarity to the types of N. antecedens; it in
fact represents one of the best documentations in the litera-
ture of the morphological variability of that species.

OCCURRENCE. South-West UK (Arn), Pennines (Asb-Arn),
Franco-Belgian Basin (Pnd-Arn), U. Silesia (Asb-Arn),
Svoge (?Arn).

Neuropteris bohdanowiczii (Zalessky) Gothan

* 1907 Sphenopteris bohdanowiczi Zalessky: 33, 65; pl. 2,
fig. 2.

8 1913 Neuropteris bohdanowiczi (Zalessky) Gothan: 210;
pl. 44, fig. 4; pl. 53, fig. 1.

REASON FOR GENERIC ASSIGNMENT. The relatively weakly
developed midvein, the slightly flexuous lateral veins, and the
tendency of the pinnules to be fused to the rachis.

COMMENTS. This is a relatively poorly known species, the
best documented records being by Gothan (1913) and Kota-
sowa (1968). Zalessky (1907) suggested that it should be
placed in a new subgenus, Sphenopteris subg. Neurospheno-
pteris. However, it is doubtful if the latter is validly published
according to the ICBN (Danzé 1956).

OCCURRENCE. U. Silesia (Alp).

Neuropteris ervedosensis (Teixeira) Wagner

* 1942 Mixoneura ervedosensis Teixeira: 8; pl. 1.
S 1963 Neuropteris ervedosensis (Teixeira) Wagner: 27.

REASON FOR GENERIC ASSIGNMENT. The apparent occurence
of enlarged, ‘forma impar’-like pinnules in the lower part of
the frond. Also, the veining, although rather less dense, has
some resemblance to that of N. ovata and N. flexuosa.

COMMENTS. Opinion is divided as to the affinities of this
species. De Jong (1974) regards it as being closely related to
N. ovata, while Wagner & Sousa (1983) state that there is
probably no relation with this species or N. flexuosa. As
stated above, its affinities seem to lean towards Neuropteris,
particularly as it probably has enlarged pinnules low in the
frond, but this must be taken as provisional, at least until
better information on the frond architecture becomes avail-
able.

OCCURRENCE. N. Portugal (WeD).

Neuropteris flexuosa Sternberg

T 1823 Osmunda gigantea, var. B Sternberg: pl.32, fig.2.

* 1825 Neuropteris flexuosa Sternberg: xvi.

T 1959 Neuropteris ovata Hoffmann, forma flexuosa (Stern-
berg) Crookall: text-fig. 52.

REASON FOR GENERIC ASSIGNMENT. Epidermal structures
(Cleal & Zodrow 1989) and close similarity of pinnule form to
N. ovata.

COMMENTS. The European records of this species, other than
those in Britain, are misidentifications of species such as
Laveineopteris tenuifolia (e.g. Novik 1952, 1954) and L.
jongmansii (e.g. Corsin 1932).

OCCURRENCE. South-West UK (WeD-Can), S Portugal
(WeD).

Neuropteris ghayei Stockmans & Williére

T 1933 Neuropteris grangeri Brongniart; Stockmans: pl.12,

fig.3.
* 1954 Neuropteris ghayei Stockmans & Williére in Pastiels
& Williére: 59.

REASON FOR GENERIC ASSIGNMENT. Underlying similarity in
pinnule form with N. heterophylla and N. obliqua. Also, on
the presence of large ‘impar’-type pinnules (Laveine 1967:
pl.49, fig.5), similar to those found in the lower part of the N.
obliqua frond.

C.J. CLEAL AND C.H. SHUTE

OCCURRENCE. Franco-Belgian Basin (Lan), NW Germany
(Lan)

Neuropteris heterophylla (Brongniart) Sternberg
15-16

* 1822 Filicites (Nevropteris) heterophyllus Brongniart: 239;
pl. 2, fig. 6.
1825 Neuropteris heterophylla (Brongniart) Sternberg:
XVi.
T 1831 Nevropteris (sic) heterophylla Brongniart: pl. 71
(neotype — vide Laveine, 1967).
T 1967 Neuropteris heterophylla Brongniart (sic); Laveine: |
pl. A. |

Figs |

REASON FOR GENERIC ASSIGNMENT. Type species.

COMMENTS. Both epidermal structures and frond architec- |
ture have been documented by Cleal & Shute (1991a).
Specimens of this species have sometimes been recorded as _
Neuropteris grangeri Brongniart (see comments by Laveine |.
1967).

The specimens from the Donets that have been illustrated |
in the literature as N. heterophylla appear to be misidentified |
specimens of Laveineopteris loshii (Novik 1952: pl. 61, figs
14; 1954: pl. 20, figs 5-6). However, Fissunenko and
Laveine (1984) claim that true N. heterophylla occurs here
and, in view of Laveine’s familiarity with the type specimens, |

the record has been accepted.

OCCURRENCE. Pennines (Duc), Franco-Belgian Basin (Lan-
Bol), NW Germany (Lan-Bol), U. Silesia (Lan-Duc), Donets
(Yea-WeD).

Neuropteris obliqua (Brongniart) Zeiller Figs 13A—B,

T 1833 Pecopteris obliqua Brongniart: pl. 96, figs 14.

* 1834 Pecopteris obliqua Brongniart: 320-321.

’ 1888b Neuropteris obliqua (Brongniart) Zeiller: 284-289.

1953a Neuropteris marginenervis Jongmans: 15; pl. 5, figs

29-30; pl. 6, figs 31-32 (vide Laveine, 1967).

.v 1959 Neuropteris lanarkiana Kidston ex Crookall: 174; pl.
50, figs 1-2.

T 1967 Neuropteris obliqua (Brongniart) Zeiller; Laveine:
pl. E, figs 1-2.

REASON FOR GENERIC ASSIGNMENT. Epidermal structure
(Cleal & Shute 1992) and frond architecture (Fig. 13; Gotha
1953: fig. 7; Scott 1978: pl. 27, fig. 1).

COMMENTS. The frond architecture of this species has still t
be fully documented, but the available evidence suggests tha
it was essentially similar to that of N. heterophylla, excep
that the pinnules at the base of the frond were rather large
(the so-called ‘forma impar’-type — e.g. Crookall (1959: pl.47
fig.4).

Crookall (1959) distinguished N. lanarkiana from N. obli
qua by its more slender, triangular and widely-spaced pin
nules, and less dense and straighter veins. However, suc
pinnules are not atypical in the N. obliqua fronds, and can b
compared with a specimen figured by Laveine (1967: pl. 50
fig. la).

Stockmans & Williére (1965: pl. 3, fig. 8; pl. 5, figs 1-2
document specimens from NW Spain as this species, an
Laveine (1967) has agreed with the identity of at least some 0

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 27

them. However, the specimens are very fragmentary and, on
their own, are inadequate for identification. There are no
better documented specimens of this species from the Iberian
Peninsula (the records by Wagner & Bowman 1983 and
Alvarez- Vazquez in Wagner 1990 are unillustrated).

OCCURRENCE. South-West UK (Lan-Duc), Pennines (Lan-
Bol), Franco-Belgian Basin (Mrd-Bol), S. Limburg (Lan-
Duc), NW Germany (Mrd-Bol), NE Germany (Yea-Bol),
Lublin (Lan-Bol), U. Silesia (Yea-Bol), Intra-Sudetic Basin
(Lan), Svoge (Lan), Donets (Yea-Bol), N. Caucasus (?Kin-
Lan), Turkey (Yea-Duc), Alps (Lan-Alps), S. Spain (Duc).

Neuropteris ovata Hoffmann Fig. 17A-B

* 1826 Neuropteris ovata Hoffmann: 266.
T 1827 Neuropteris ovata Hoffmann: pl. 1b, fig. 6 (vide
Satzwedel, 1969).
1888a Nevropteris stipulata Zeiller: 255; pl. 29 fig. 5 (tenta-
tively suggested by Wagner & Alvarez-Vazquez,
1991).
._p 1960 Neuropteris valdensis (Heer) Jongmans: 57; pl. 18,
in. JOFe
'T 1969 Imparipteris ovata (Hoffmann) Gothan; Saltzwedel:
pl. 24 figs 1-2.
-.y 1973 Mixoneura polyneura Doubinger & Germer: 50-51;
pl. 1, fig. 2.
.) 1974 Neuropteris germeri de Jong: 58; pls 21-22.
_.v 1975a Neuropteris pilosa Doubinger & Germer: 18; pl. 7,
fig. 1.

REASON FOR GENERIC ASSIGNMENT. Epidermal structures
| (Barthel 1962, Cleal 1985, Cleal & Zodrow 1989) and frond
architecture (Zodrow & Cleal 1988).

COMMENTS. Cleal & Zodrow (1990) recognize varieties of
this species, based partly on differences in epidermal struc-
ture. Also, Wagner (1963) assigns most of the Stephanian
‘examples of the species to a separate variety (var. grandeuryi
' Wagner), based on the lateral veins being denser and less
‘oblique to the pinnule margin. While recognizing that these
| Varieties almost certainly have some biological validity, they
will not be separated in this analysis.

The identity of upper Stephanian C specimens from north
Portugal, described by Wagner & Sousa (1983) as Neuropteris
ovata var. pseudovata Gothan & Sze, centers on one of the
most contentious issues concerning the taxonomy of this
species, viz. the difference between it and Neurocallipteris
neuropteroides. There have been many analyses of this prob-
‘lem, the most detailed being by Zalessky (1909), Barthel
(1976), Setlik (1980) and Wagner & Sousa (1983). Setlik and
Wagner & Sousa have shown there are certain very subtle
differences in pinnule form, but without the evidence of
cuticles it is far from certain that they would be regarded as
Sufficient justification for separating them as species, let
alone in different form-genera. It would seem that two quite
_\separate groups of trigonocarpaleans have developed analo-
gously similar pinnule morphologies, perhaps in response to
similar environmental pressures. Consequently, identifying
specimens with this type of pinnule in the upper Stephanian,
f epidermal characters are unknown, is very difficult, if not
mpossible. Wagner & Sousa’s Portugese specimens have
some of the characters (somewhat subtriangular pinnules,
veins slightly oblique to pinnule margin) that tend to be
commoner in N. neuropteroides than N. ovata. We have

therefore provisionally transferred their specimens to N.
neuropteroides. However, we recognize that this needs to be
confirmed by epidermal evidence or, if this is impractical, by
larger specimens showing the form of the intercalated pin-
nules (cf. Setlik 1980).

M. polyneura and N. pilosa were separated from N. ovata
on minor characters of venation and surface detail, and their
distinction cannot be maintained (Cleal 1985). N. germeri,
which was described mainly on just two specimens, has a
veining pattern and pinnule shape compatible with the larger
forms of N. ovata, especially those of the var. sarana as
described by Cleal & Zodrow (1989). The veining density is a
little lower than is typical, but can probably be accommo-
dated within the lower end of the range of variation. As the
types of N. germeri are associated with more typical frag-
ments of N. ovata var. sarana (Bertrand) Cleal & Zodrow,
there seems little reason for distinguishing them taxonom-
ically.

The type of N. stipulata is strikingly similar to N. ovata in
pinnule shape, being relatively squat, having a basiscopic
auricle and a short midvein. If Zeiller’s drawing of the type of
N. stipulata is accurate, the vein density is c.40 veins per cm,
which is compatible with the form of N. ovata, normally
found in the Stephanian, and known as var. grandeuryi
Wagner.

OCCURRENCE. South-West UK (WeD-Can), Pennines
(WeD), Franco-Belgian Basin, (WeD), NW Germany
(WeD), U. Silesia (WeD), Saar-Lorraine (WeD), Donets
(?Bol-?Bar), N. Caucasus (WeD, StB), Turkey (WeD), Alps
(WeD-StC), Massif Central (Bar-StB, ?StC), NW Spain
(WeD-StB), S. Spain (StC).

Neuropteris parvifolia Stockmans

1933 Neuropteris parvifolia Stockmans: 28-29, pl. 8, figs
1-5.

REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule
morphology and underlying venation pattern to N. obliqua
(see Laveine 1967).

OCCURRENCE. Franco-Belgian Basin (Duc-Bol), NW Ger-
many (Duc-Bol), NE Germany (Duc-Bol), Lublin (Duc-
Bol), Alps (Bol).

Neuropteris plicata Sternberg

1833 Neuropteris plicata Sternberg: 70; pl. 19, figs 1,3.

REASON FOR GENERIC ASSIGNMENT. Close similarity of the
pinnules to N. ovata.

COMMENTS. This is so similar to N. ovata that it is far from
certain that it is a distinct species. Although Setlik (1921)
attempted an analysis of the morphological variation of the
pinnules, the results were equivocal on this point. If they are
the same species, then Sternberg’s species would take prior-
ity. In view of the important palaeobotanical and biostrati-
graphical role played by N. ovata, a more thorough analysis
of N. plicata should be undertaken before any nomenclatural
changes are proposed.

OCCURRENCE. Intra-Sudetic Basin (WeD).

Neuropteris praeovata (Némejc) Cleal & Shute, comb.
nov.

* 1949 Mixoneura praeovata Némejc: 17-18; text fig. 2; pl.
4, figs 1-7.

REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule
shape and venation to N. ovata.

COMMENTS. As pointed out by Némeje, this clearly belongs
to the group of neuropterids allied to N. ovata, from which it
can be reliably distinguished only by the more prominent
midvein. Also, it is the only known member of this group of
species to occur below the Westphalian D.

OCCURRENCE. Intra-Sudetic Basin (Bol).

Neuropteris resobae Cleal
* 1981 Neuropteris resobae Cleal: 79, pls 1-2.

REASON FOR GENERIC ASSIGNMENT. Dense venation with
relatively weak midvein. Pinnule limb often partly fused to
rachis, and with basiscopic auricle.

COMMENTS. Little is known of the frond architecture, other
than that the ultimate pinnae are terminated by a single apical
pinnule, and nothing of the epidermal structures. Its reten-
tion in this form-genus is thus based entirely on pinnule
morphology and venation, which clearly cannot be conclu-
sive. The only other (albeit negative) piece of evidence is
that, despite the size of the collection on which the species
was described, not a single cyclopterid pinnule was found in
association.

OCCURRENCE. NW Spain (Duc-WeD).

Neuropteris schaeferi Doubinger & Germer

* 1975a Neuropteris schaeferi Doubinger & Germer: 10-11;
pl. 4, fig. 1.

REASON FOR GENERIC ASSIGNMENT. Similarity of epidermal
structure to N. ovata (see Saltzwedel 1968).

COMMENTS. This species is not really adequately docu-
mented, as only one small specimen has been described in the
literature. However, in view of the epidermal evidence
presented by Saltzwedel (1968), it has been included in the
present analysis.

OCCURRENCE. Saar-Lorraine (StB).

Neuropteris semireticulata Josten

* 1962 Neuropteris semireticulata Josten: 39-40; pl.3, figs
2-5.

REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule
morphology and underlying venation pattern to N. obliqua
(see Josten 1962).

OCCURRENCE. South-West UK (Duc-Bol), Pennines (Duc),
Franco-Belgian Basin (Duc-Bol), NW Germany (Duc-Bol),
Lublin (Bol). i

Neuropteris willierei Laveine

* 1967 Neuropteris willierei Laveine: 224-227; pl.62.

C.J. CLEAL AND C.H. SHUTE |

REASON FOR GENERIC ASSIGNMENT. It is almost indistin-
guishable from N. parvifolia, differing only in having smaller,
more broadly attached pinnules, and occurring stratigraphi-
cally lower (Laveine 1967).

OCCURRENCE. Franco-Belgian Basin (Lan-Duc), NW Ger-
many (Lan-Duc).

Form-genus PARIPTERIS Gothan

Paripteris gigantea (Sternberg) Gothan

* 1821 Osmunda gigantea Sternberg: 33; pl.22.

‘1941 Paripteris gigantea (Sternberg) Gothan: 427.

T 1953 Neuropteris gigantea Sternberg; Havlena: pl. 4; pl.
5), he, 2.

.v 1959 Neuropteris maltbyensis Crookall: 164; pl. 33, figs
7-8.

? 1965 Paripteris veeni Stockmans & Williére: pl. 2, figs 4-7
(vide Wagner & Bowman 1983).

REASON FOR GENERIC ASSIGNMENT. Type species.

OCCURRENCE. South-West UK (Lan-Duc), Pennines (Lan-
Duc), Franco-Belgian Basin, (Mrd-Duc), NW Germany
(Mrd-Bol), NE Germany (Kin-Lan), Lublin (Kin-Duc),
Intra-Sudetic Basin (Lan-Bol), U. Silesia (Mrd-Bol), Svoge
(Yea-Lan), Donets (Kin-Duc), N. Caucasus (?Kin-Lan),
Turkey (Lan), Alps (Lan), Pyrenees (Kin), NW Spain (Mrd,
Lan), S. Spain (Lan).

Paripteris linguaefolia (Bertrand) Laveine

* 1930 Neuropteris linguaefolia Bertrand: 31-32; pl. 15.
’ 1967 Paripteris linguaefolia (Bertrand) Laveine: 266-267;
pls 77-78.

REASON FOR GENERIC ASSIGNMENT. Frond architecture.

OCCURRENCE. Franco-Belgian Basin (Duc-Bol), NW Ger-
many, (Bol), Lublin (Duc-Bol), Intra-Sudetic Basin (Duc-
Bol), Saar-Lorraine (Duc-Bol), Donets (Duc-WeD), Turkey
(Duc), Alps (Duc-Bol), NW Spain (Duc), S. Spain (Duc).

Paripteris linguaenova (Bertrand) Cleal & Shute,
comb. nov.

* 1930 Neuropteris linguaenova Bertrand: 29; pls 13-14.

eee
— me tad

ie er fe, ee ,

REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule
shape and venation to P. linguaefolia.

COMMENTS. Laveine (1967) assigned the types of this species
to P. pseudogigantea. However, they are significantly larger
(many are longer than 4 cm), have a thinner midvein, and
lateral veins that diverge from the midvein at a narrower
angle. More problematic is its distinction from P. linguae-
folia. Bertrand (1930) separated them because P. linguaefolia
has virtually no midvein, and ‘la disposition et l’aspect des
nervures sont trés différents. . .". As the types of P. ling-
uaenova occur within the stratigraphical range of P. linguae-
folia, these differences really need to be re-examined more
critically. For the time being, however, the separation has
been retained.

OCCURRENCE. Saar-Lorraine (Bol).

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 29

Paripteris pseudogigantea (Potonié) Gothan Fig. 13C-D

* 1897 Neuropteris pseudogigantea Potonié: 113; text
fig. 102.
1941 Neuropteris scheuchzeri forma minor Novik: 457; pl.
Dips 5 7h
* 1953 Paripteris pseudogigantea (Potonié) Gothan: 63-64;
pl. 37, figs 1-4.

_ REASON FOR GENERIC ASSIGNMENT. Frond architecture
(Laveine 1967: pls 73-76).

:
/ OCCURRENCE. South-West UK (Duc-Bol), Pennines (Duc-
_ Bol), Franco-Belgian Basin (Duc-Bol), NW Germany (Duc-

Bol), NE Germany (Duc-Bol), Lublin (Duc-Bol), Saar-
| Lorraine (Duc-Bol), Donets (Duc-Bol), NW Spain (Duc).

Paripteris schuetzei (Potonié) Daber
:

| * 1903 Neuropteris schiitzei Potonié: 399.

T 1912 Neuropteris schiitzei Potonié: 122; fig. 84.

_* 1963b Paripteris schiitzei (Potonié) Daber: 1212, fig. 2.
.

| REASON FOR GENERIC ASSIGNMENT. Frond architecture and
» general aspect of pinnules.

} OCCURRENCE. S. Limburg (Lan), U. Silesia (Duc).

Form-genus SPHENONEUROPTERIS Shchegolev

* Sphenoneuropteris brongniartii Shchegolev

)

1979 Sphenoneuropteris brongniartii Shchegolev: 159; pl.
53, fig. 2.

| REASON FOR GENERIC ASSIGNMENT. General similarity in
| -pinnule morphology and venation with type species.

_ COMMENTS. The distinction between this and S. elegans is far
) from clear, and there must be a strong likelihood that they
. are synonyms.

| OCCURRENCE. N. Caucasus (StC).

| Sphenoneuropteris dimorpha (Lesquereux) Cleal &
Shute, comb. nov.
IT 1879 Pseudopecopteris dimorpha Lesquereux: pl. 35, figs
1-6.
1880 Pseudopecopteris dimorpha Lesquereux: 201.
1978 Neuropteris dimorpha (Lesquereux) Boersma: 59;
pl 8, tig. 3; pl. 12; figs 1-6.

REASON FOR GENERIC ASSIGNMENT. Mainly the venation
(widely forking veins, oblique to pinnule margin, producing
low vein density), and the large, lax-limbed pinnules.

COMMENTS. Little is known of the frond architecture and
nothing of the epidermal structure of this species. However,
the pinnules show a remarkable similarity, especially in their
venation (e.g. Doubinger & Germer 1975b, pl. 4), to the
ypes of Sphenoneuropteris.

This species is often thought to have characteristically
leeply-lobed pinnules (e.g. Wagner 1958). However,
30ersma (1978) showed that this was at least partially a
onsequence of the thin limb of the pinnules, which rarely lay
lat in the matrix, and would undulate in and out of the plane
long which the fossil was split. It is unlikely that this can

explain all specimens with undulate margins, but the remnant
examples may simply be from the distal regions of pinnae,
where pinnules are in transition to ultimate pinnae.

This species was initially assigned to Pseudopecopteris
Lesquereux, 1880. However, this form-genus was not typified
and included within it was a variety of disperate types of
frond; it is thus a nomen dubium, and cannot be used as an
alternative name for Sphenoneuropteris.

OCCURRENCE. Saar-Lorraine (Bol-StB).

Sphenoneuropteris elegans Shchegolev

1979 Sphenoneuropteris elegans Shchegolev: 158; pl. 54,
figs 1,2.

REASON FOR GENERIC ASSIGNMENT. Type species.

OCCURRENCE. N. Caucasus (StC).

Sphenoneuropteris nemejciana (Purkynova) Cleal &
Shute, comb. nov.

1971 Neuropteris nemejciana Purkynova: 168; pls 10-11.
1971 Neuropteris venceslai Purkynova: 171; pl. 12.

REASON FOR GENERIC ASSIGNMENT. Similarity of pinnule
form and venation to S. dimorpha.

COMMENTS. Purkynova’s specimens occur stratigraphically
lower than any of the other species included in Spheno-
neuropteris. However, it has many of the characteristic gross
morphological features of that form-genus, including large
pinnules (30-35 mm long) with a lax limb and somewhat
undulate margin, and a low vein density (16 veins per cm on
pinnule margin).

The type and only known specimen of N. venceslai ori-
ginated from the same locality and horizon as the types of S.
nemejciana. It has similarly large, relatively thin-limbed pin-
nules, thin midvein, and low vein density; compare for
instance the specimen figured by Purkyfova on her pl. 11, fig.
2. The pinnules have a more acute apex, and are marginally
larger, but not execssively so; the largest recorded pinnule of
S. nemejciana is 5.5 cm long, as opposed to 7.5 cm in N.
venceslat. All in all, there seems little reason to regard these
as separate species.

OCCURRENCE. U. Silesia (Lan).

Sphenoneuropteris praedentata (Gothan) Cleal &
Shute, comb. nov.

* 1909 Neuropteris praedentata Gothan: figs 1,2.

REASON FOR GENERIC ASSIGNMENT. Based mainly on vena-
tion (widely forking veins, oblique to pinnule margin, pro-
ducing low vein density), and the relatively large,
subtriangular pinnules.

COMMENTS. The general aspect of the pinnules, particularly
the venation, seems to exclude this from Neuropteris as it is
interpreted in this work. The venation seems to fit in far
better with that given in the diagnosis of Sphenoneuropteris
given by Shchegolev (1979). It is recognized that this is far
from a satisfactory basis for recognizing ‘natural’ form-
genera. However, until cuticle and frond architecture data
become available, Shchegolev’s form-genus provides a conve-
nient repository for this species.

The only large specimens of this species to have been
published are in Zeiller (1888a: pl. 26) and Zeiller (1906: pl.
26), both under the name Neuropteris crenulata Brongniart.
They both show bipinnate frond fragments, with intercalated
pinnules on the penultimate rachis. Laveine (1967: text-fig.
6d) interprets the 1906 specimen as essentially a pinnate
frond. However, the penultimate rachis in the 1888 specimen
is noticeably curved, suggesting that it might be from a
bipartite frond, similar to that present in many of the other
neuropteroid form-genera.

The numerous records of this species from the Iberian
Peninsula have been analysed by Knight (1983). He has
concluded that, although they show some similarity to S.
praedentata, they differ in having smaller, thinner-limbed
pinnules with weaker crenulations on the margin, and thinner
veins. They have since been transferred to a separate species,
S. wagneri (see below). Significantly, Knight also observed
that the Spanish material shared some features in common
with S. dimorpha, providing some support for the idea that S.
dimorpha, S. praedentata and S. wagneri cluster together to
form a reasonably natural form-genus.

OCCURRENCE. (?)Saar-Lorraine (Bar), Massif Central (Bar-
St@)?

Sphenoneuropteris wagneri (Lorenzo) Cleal & Shute,
comb. nov.

* 1980 Mixoneura wagneri Lorenzo: 11-13; pl. 1.

REASON FOR GENERIC ASSIGNMENT. The large, relatively lax
pinnules with a wide venation.

COMMENTS. This species was established for the Spanish
specimens that were traditionally assigned to “Neuropteris’
praedentata (see comments on previous species).

OCCURRENCE. NW Spain (Bar-Aut).

Species of uncertain taxonomic position

Included here are those species which, although clearly
circumscribed and thus ‘good’, cannot be readily assigned to
any of the above form-genera. Cuticular evidence is lacking,
and their pinnule and pinna morphologies do not provide any
obvious comparison with one or other of the more completely
known species.

Neuropteris bourozii Laveine
* 1967 Neuropteris bourozii Laveine: 152; pls 23-25.

COMMENTS. Some of the pinnules of this species show simil-
arities to Laveineopteris (Laveine 1967: pl.24, fig.5), while
others are of a more typical neuropterid-type (Ibid. pl.23,
fig.5). Laveine (1967) assigned specimens from the Pennines
Basin figured by Bolton (1926: pl. 6) to this species, but they
almost certainly belong to L. tenuifolia.

OCCURRENCE. Franco-Belgian Basin (Duc), NW Germany
(Duc).

Neuropteris cordata Brongniart

1831 Neuropteris cordata Brongniart: 229; pl. 64.
1890 Nevropteris Raymondii Zeiller: 147; pl. 9a, fig. 4.

C.J. CLEAL AND C.H. SHUTE

1893 Neuropteris pseudoblissii Potonié: 137.
1964 Mixoneura raymondii (Zeiller) Wagner: 9.

COMMENTS. This species cannot readily be fitted into any of
the other form-genera. The general aspect of the pinnules
suggests affinities with Neurocallipteris or possibly even Neu-
ropteris, but what little is known of the frond architecture
(e.g. Langiaux 1984: fig. 111) would seem to separate it from
both genera. The species is in clear need of a revision.

The type and only known specimen of N. raymondii Zeiller
(Mixoneura raymondii (Zeiller) Wagner) was figured photo-
graphically by Doubinger (1956: pl. 12, fig. 3; pl. 13, fig. 1). It
originated from the Mont Pel Formation in the Autun-Epinac
Basin, and occurs together with specimens of N. cordata. The
pinnules are rather smaller (c.16 mm long) than is typical for
N. cordata but the venation is very similar. Doubinger (1956)
claims that the veining density is higher in N. raymondii, but
the measured value of 22 veins per cm on the pinnule margin |
is quite compatible with some of the smaller forms of N.
cordata (cf. Zeiller 1906: pl. 27, fig. 3). In view of the |
evidence of association and of the similarity of the venation,
it seems reasonable to assume that N. raymondii is merely a |
small-pinnuled form of N. cordata, possibly from the more
distal regions of the frond.

N. pseudoblissii is still being recorded in the modern
literature for specimens from the upper Stephanian with very
elongate pinnules, but which are otherwise very close to N.
cordata. Zeiller (1888a), who figured the types of Potonié’s
species under the incorrect name Neuropteris blissii, noted
that isolated fragments would be difficult to distinguish, and
it is also significant that the two species almost invariably
occur together (e.g. see records in Doubinger 1956). There
thus seems little justification for separating the two species.

The single specimen from the Duckmantian of the Pen-
nines figured by Crookall (1959: pl. 41, fig. 5) as N. pseudo-
blissii is an indeterminable fragment, possibly of a
mariopterid.

OCCURRENCE. Massif Central (StB-StC), Pyrenees (StC),
Alps (StC), N. Portugal (StC).

Neuropteris duprei Laveine
* 1967 Neuropteris duprei Laveine: 164, pl.29, figs 14.

COMMENTS. This is a very distinctive species with elongate,
often asymmetrical pinnules and very oblique lateral veins.
The only other similar material reported from Europe are th
specimens described by Némejc (1949, pl.1, figs 1-8) “4
Odontopteris stradonicensis Andra. There is also a recor
from the Langsettian of NW Spain (Wagner & bona
1983), but it is not illustrated.

OCCURRENCE. Franco-Belgian Basin (Lan-Duc).

Neuropteris dussartii Laveine
* 1967 Neuropteris dussartii Laveine: 191; pl.48.

COMMENTS. Laveine argued that this species shared a num
ber of features in common with laveineopterid species such a
L. loshii and L. rarinervis. However, Laveine also pointe
out certain similarities with Neuropteris ovata, such as th
presence of a basiscopic auricle on some of the pinnules.

OCCURRENCE. South-West UK (Bol-WeD), Franco-Belgia
Basin (WeD), NW Germany (WeD).

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 31

Neuropteris teberdensis Shchegolev

* 1979 Neuropteris teberdensis Shchegolev: 163; pl. 51; nl.
Sa ike, WS pol, S3i5 104, Il,

COMMENTS. The pinnules of this very late species show a
_ marked resemblance to Neuropteris ovata, suggesting that it

is a true neuropterid. However, one of the specimens (Shche-

golev 1979: pl. 52, fig. 1) suggests that the frond might have

been only bipinnately divided, with intercalated pinnules on

the primary rachis branches. This fact, together with its high
Stratigraphical position, suggests that the species may instead
- belong to Neurodontopteris.

_ OCCURRENCE. N. Caucasus (StC).

Neuropteris zeilleri de Lima

_T 1864 Neuropteris cordata Brongniart; Géppert: 100; pl.
11, figs 1-2.
* 1890 Neuropteris zeilleri de Lima: 140.

COMMENTS. This species has been widely quoted in the
literature as occurring in the upper Stephanian of Europe
(e.g. Havlena 1953, Doubinger 1956, Wagner 1963, Vetter
1968, Wagner & Sousa 1983). As pointed out by Zeiller
/(1906) and Vetter (1968), however, there are problems with
ithe typification of the species; that quoted above is the one
‘normally accepted, but it is far from clear if de Lima regarded
Goppert’s specimens or his own Portugese specimens as
types. The distinction from Neuropteris cordata is also far
‘from clear and according to Zeiller is based mainly on the fact
that there is not a single midvein, but a number of separate,
fine veins lying along the long axis of the pinnules. This
distinction has never been properly documented and there
must be a strong suspicion that it is purely taphonomic.
Whatever the outcome, however, there can be little doubt
hat N. zeilleri will end up in the same form-genus as N.
vordata, whatever that will prove to be (see above).

JCCURRENCE. (?) Intra-Sudetic Basin (Aut), Massif Central
2StB, StC-Aut), NW Spain (StB, ?StC), N. Portugal (StC-
Aut).

Nomina dubia

he first group of species included here were initially
escribed on just one or two fragments and additional
jaterial has not been published. There is thus insufficient
vidence of morphological variation to be able to recognize
ie species reliably, or of features such as frond architecture
‘epidermal structure, by which their generic position could
= assertained. They are listed below without further com-
ent.

"paripteris flabellinervis Gothan, 1953: 59; pl. 9, figs 2-3; pl.
28, fig. 2; pl. 30, fig. 6.

europteris asturiana Jongmans MS ex Wagner, 1962: 757
{nomen nudum].

2uropteris beveridgei Crookall, 1959: 189, pl. 40, fig. 4.
puropteris bulupalganensis Zalessky in Zalessky &
Chirkova, 1933: 9; fig. 1.

?uropteris(?) delasii Zeiller, 1892: 45; pl. 8, fig. 6.
"uropteris dispar Zeiller, 1888a: 253; pl. 29, fig. 6.
‘uropteris horrida Zeiller, 1888a: 251; pl. 32, figs 1-2.

|
|

Neuropteris jugosa Kidston ex Crookall, 1959: 164; pl. 41, fig.
3

Neuropteris matheronii Zeiller, 1888a: 245; pl. 28, fig. 7.

Neuropteris pseudoimpar Stockmans & Williére, 1953: 235:
pl. 44, fig. 2; pl. 50, fig. 12.

Neuropteris squarrosaeformis Kidston ex Crookall, 1959: 163:
pl.50, fig.6.

Neuropteris subsessilis Stockmans & Williére, 1955: 14; pl. 8,
fig. 1.

Neuropteris waltonii Stockmans & Williére, 1953: 227-228.

In addition to the above, there are a number of other species,

for which more specimens are known, but which are still

impossible at present to identify reliably. These require

further comment.

Mixoneura muensterifolia Némejc

1949 Mixoneura muensterifolia Némejc: 15-16; pl. 3, figs
10-14.
1949 Mixoneura grandifolia Némejc: 18-20; text fig. 4.

COMMENTS. Némejc established this species for a number of
fragments from the middle Westphalian, that were claimed to
have a pinnule shape similar to Neuropteris obliqua, but with
more flexuous veins. These are similar to the characters used
to define N. semireticulata, of which it would be an earlier
synonym. However, the illustrations used by Némeje are
poor and the specimens fragmentary. It would thus be unwise
to give it priority over N. semireticulata, at least until Ném-
ejc’s species is better documented.

Némejec reported larger pinnules in close association with
M. muensterifolia, and used them as the types of another new
species, M. grandifolia. However, the figured specimens
would seem to correspond with forma impar-type pinnules
found in the proximal parts of the fronds of the N. obliqua
group. It is thus almost certain that they are conspecific with
the specimens that he assigned to M. muensterifolia.

Neuropteris arberi Crookall

* 1959 Neuropteris arberi Crookall: 148; pl. 50, fig. 7; pl.
51, figs 2-4.

COMMENTS. Based on three fragments, none of which show
details of the apical pinnules or the pattern of lobing. Their
affinities may be more mariopterid than neuropterid. Remy
& Remy (1975) attempted to use this species for German
specimens, but it is difficult to see how this can be justified in
the light of the extremely imperfect types.

Neuropteris kosmannii Potonié

* 1903 Neuropteris kosmanni Potonié: 399.

T 1913 Neuropteris kosmanni Potonié; Gothan: pl. 47, fig.
3; pl. 50, figs 14.

COMMENTS. Although this species periodically re-appears in
the literature (e.g. Kotasowa 1968), it has only ever been
described from small fragments. They all show vaulted,
extremely thick-limbed pinnules, often with a somewhat
undulate margin, quite atypical for any of the neuropteroid
form-genera, with the possible exception of Margaritopteris.
In the absence of more complete material, it is impossible
either to give it a useful circumscription as a species, or to
determine their generic position.

Neuropteris lubnensis Havlena

* 1953 Neuropteris lubnensis Havlena: 153-154; pl. 6, figs
1-2.

COMMENTS. This is based on forty-eight specimens preserved
in a sandstone, although only two were figured. They are
undoubtedly unusual, having very large pinnules (up to 4 cm
long and 2 cm wide), and do not fit into any previously
described species. However, the arenaceous matrix is far
from perfect for preserving this type of fossil, and the number
of specimens illustrated is inadequate to determine the range
of morphological variation, let alone frond architecture.
Much better material needs to be documented before any-
thing can be done with this species.

Neuropteris montana Heer

* 1879 Neuropteris montana Heer: 22; pl. 6, figs 22, 23.
T 1960 Neuropteris montana Heer; Jongmans: pl. 21, fig.
121.

COMMENTS. From the form of the distal part of the pinna, the
type clearly belongs to a paripinnate frond, presumably of the
Potonieaceae. It is reputed to originate from the Cantabrian
or lower Barruelian, which is far higher stratigraphically than
Paripteris normally occurs. The venation is very poorly
preserved, but may be anastomosed with very elongate
vein-meshes. If so, then it may belong to Linopteris neu-
ropteroides (Gutbier) Potonié, 1899, which sometimes occurs
as high as Barruelian. However, the material is really inad-
equate to give an unequivocal statement on this.

ROBUSTNESS OF GENERIC TAXONOMY

The statistics of this taxonomic analysis are summarized in
Table 2 and Fig. 18. A total of 101 neuropteroid species have
been recorded from Europe over the last half century, of
which nearly a half (43-5%) are either unsatisfactory because
they are based on insufficient material, or are later synonyms
of other species. Of the remaining fifty-seven ‘good’ species,
all but six (10-5%) can be assigned to one or other of the 9
form-genera summarized in the early part of this paper.

Total good species (0.6)

Inedequately circumscribed (0.2)

Synonyms (0.3)

Fig. 18 The robustness of the taxonomy of neuropteroid fronds. (a) The proportions of synonyms, inadequately described and ‘good’ species
among all those neuropteroids used since 1940. (b) The proportion of the ‘good’ species belonging to each of the form-genera.

Neurodontopteris (0.0)

C.J. CLEAL AND C.H. SHUTE

The six species that cannot yet be placed in our generic
classification fall into three groups.

1. N. cordata and N. zeilleri appear closely related to each
other and it is far from certain that they are not in fact
conspecific. Although widely recorded from the Stephanian
and Autunian of France and the Iberian Peninsula, little is
known of the frond architecture and nothing of the cuticles
(the cuticles assigned to N. cordata by Barthel, 1976, in fact
belong to Neurodontopteris auriculata — see above).

2. N. duprei has unusual, asymmetrical pinnules unlike |
any of the other species included in this analysis; in fact they |
are different from any type of foliage previously assigned to |
the trigonocarpaleans. It may well belong to a new form-
genus, but details of the frond architecture and/or cuticles will
be needed before any decision on this can be made.

3. From the general aspect of the pinnules, it is likely that
N. bourozii, N. dussartii and N. teberdensis belong to either |
Neuropteris, Laveineopteris or Neurocallipteris. Again, evi- |
dence of frond architecture and/or cuticles will be needy

|
|
|

before a decision can be made on their classification.
In conclusion, the analysis has allowed us to see where the
main gaps are in our knowledge of these fossil fronds. In
particular, the 6 species that cannot currently be assigned
need to be further investigated. Nevertheless, we believe that |
the results support the essential robustness of our generic}
classification of neuropteroid fronds, and points to it being a|
potentially useful tool for understanding more clearly the
distribution of these plants. |

DIVERSITY ANALYSIS

Diversity of the neuropteroids as a whole

As a by-product of this study, whose original goal was merely
to ascertain the robustness of the generic classification of
neuropteroid fronds, we have built up a database of the
stratigraphical and geographical distribution of species within
Europe. This would appear to invite further analysis of
diversity variations. Diversity analysis has become a popular
pursuit in recent years, but can be prone to serious problems|

Paripteris (0.1)

Sphenoneuropteris (0.1) |

Neuropteris (0 3)
Uncertain affinities (0.1)

Laveineopteris (0.2)

Neurocallipteris (0.1)

Macroneuropteris (0.1)
Margaritopteris (0.0)

Neuralethopteris (0.1)

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 33

Table 2 Statistics of neuropteroid taxonomy

No. of
Form-genera species % (Total) % (Good)
Laveineopteris 9 8-91% 15-79%
Macroneuropteris 4 3-96% 7:02%
Margaritopteris 1 0-99% 1-75%
Neuralethopteris 7 6:93% 12:28%
Neurocallipteris 8) 2:97% 5:26%
Neurodontopteris 1 0:99% 1:75%
Neuropteris 15 14-85% 26-32%
Paripteris 5 4-95% 8:77%
Sphenoneuropteris 6 5-94% 10-53%
- Uncertain affinities 6 5-94% 10-53%
Inadequately circumscribed 17 16-83%
Synonyms 27 26-73%
Total ‘good’ species / 56-44%
' Total ‘bad’ species 44 43-56%
Grand total 101 100-00%

) due at least in part to the tendency to use data trawled
' uncritically from the literature (cf. comments by Cleal 1988).
Our database, although based only on a small range of
species, at least has the merit of having been critically
. compiled.

| To this end, a tabulated set of statistics has been compiled
to represent diversity, first-appearances and extinctions for
each stage (Table 3). This has been done separately for each
‘of the form-genera, as well as for the group as a whole
| (including those species unassignable to any of the form-
genera).

| The diversity of the group as a whole follows a fairly simple
_pattern, showing a marked peak in the Westphalian, followed
)by a rapid decline and then a subsidiary peak in the upper
Stephanian (Fig. 19A). The Westphalian peak would seem to
be confirmed by observations made by Boulter et al. (1988)
on diversity changes in the wider plant adpression record for
ithe palaeoequatorial belt. It almost certainly reflects varia-
tions in the available non-marine strata in Europe; Niklas et
al. (1980, p. 29) demonstrated that 98:5% of plant fossil
diversity (at least between the Carboniferous and Jurassic)
can be accounted for by this single factor. Numerical data on
the available strata in each stage are not available for Europe.
However, our observations would seem to confirm the gen-
eral impression that delta-plain, fluvio-lacustrine deposits,
which presumably reflect the habitats favoured by the plants
yroducing these fronds, are at a maximum in the Westphalian
ind upper Stephanian, with a low in the Cantabrian and, to
{in extent, the Baruellian.

| Figs 19B and 19C show the patterns of appearances and
pxtinctions per stage, both corrected for variation in the
‘ength of the stage. These show curves with a similar double-
eaked form to the diversity curve. A broad correlation
‘etween species turn-over and diversity is not surprising.
fowever, the species profit/loss curve (Fig. 19D) shows a
hore interesting pattern. Up to the Kinderscoutian the
\ituation is relatively stable, but at higher stratigraphical
pvels there are major fluctuations. Peaks occur in the
sinderscoutian, Langsettian and Baruellian/Stephanian B.
‘he first of these can be correlated with the first appearance
f large-scale deltas across northern Europe; the second the

|
|

proliferation of coal-swamp conditions on the delta-tops; and
the third the expansion of intra-montane basins in central and
southern Europe. The trough in the Cantabrian presumably
reflects the change-over from predominantly paralic to pre-
dominantly intra-montane conditions over much of Europe.

So, the diversity of the neuropteroids as a whole is merely a
function of the general diversity of the tropical swamp
vegetation. If the form-genera outlined earlier in this paper
have any basis in the genetic relationships of the parent
plants, diversity patterns of the individual form-genera may
tell a different story. Fig. 20 shows the diversity curves of six
of the most abundant of the form-genera plotted separately.
This clearly shows that the story is far more complex.
However, the style of analysis dealt with so far in this paper is
not really suitable for uncovering the more detailed distribu-
tional patterns. For this, we need to look at the detailed
variations in diversity of the species within each of the
form-genera.

The problem here is the limited amount of suitable data
available. There have been studies documenting the quantita-
tive stratigraphical variations of different species, such as by
Davies (1929). However, such work is mostly old, largely
unillustrated and uses unreliable taxonomy. Also, as pointed
out by Scott (1985), there are serious weaknesses with the
sampling that was usually employed. Scott himself suggested
that quadrat analysis, similar to that sometimes used to study
living plant ecology, could produce more reliable results.
However, while quadrat analysis might prove valuable in the
detailed relationship between facies and plant fossils at a
specific locality, it would need a considerable number of such
studies before it would reveal any meaningful stratigraphical
patterns of plant fossil distribution.

We have instead adopted an alternative approach, by
looking at the numbers of localities from where a species is
recorded at different stratigraphical levels. To do this, it was
decided to restrict the analysis to one particular area, which
would help minimize potential palaeolatitudinal variations.
The area should have numerous records spread over a
reasonably long stratigraphical range. The data should also
preferably be based on identifications made by a single
authoritative palaeobotanist, thus minimizing the potential
for subjective variations in identification. In fact, only one
area was found to have all these virtues, namely the Franco-
Belgian Basin, through the monographic study by Laveine
(1967).

Species diversity analysis (Franco-Belgian Basin)

Laveine’s (1967) monograph provides a taxonomically reli-
able record of most of the neuropteroid species found in the
paralic belt between the Kinderscoutian and Westphalian D.
For each species, he individually lists the localities where they
are found in the Nord-Pas-de-Calais Coalfield, divided strati-
graphically into lower, middle and upper divisions of the
formations there. Using this data, we have plotted the
diversity curves for each species of four of the form-genera
(Figs 21-23).

Neuropteris (Fig. 21). These 7 species appear to fall into
two groups. The early group consists of N. obliqua, N.
heterophylla, N. ghayei and N. willieri, which occur predomi-
nantly in the Langsettian and basal Duckmantian (in Bel-
gium, N. obliqua is reported to extend down to the
Marsdenian, but the French records on which the present

C.J. CLEAL AND C.H. SHUTE

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> Laveineopteris + Neuralethopteris ~™* Neurocallipteris

-* Neuropteris % Paripteris 4 Sphenoneuropteris
Fig. 20 Stratigraphical diversity of six of the more abundant
neuropteroid form-genera, showing complexity hidden by

generalized graph in Figure 19a.

analysis is based only show it as far back as the Langsettian).
It then undergoes a significant decline in the lower Duckman-
tian. Only one of the species extends much beyond the
middle Duckmantian, and that is what is referred to in the
chart as the N. obliqua group. This pattern seems to be in
general agreement with what is seen in areas other than the
Franco-Belgian Basin.

As with the other neuropterids, N. obliqua sensu stricto
undergoes a marked decline in the lower Duckmantian.
However, in the upper Duckmantian there is the start of a
progressive change in the venation, which becomes more
flexuous and eventually culminates in the anastomosed-
veined form known as Reticulopteris Gothan. Details of this
gradual change in venation through the Duckmantian and
Bolsovian have been documented by Josten (1962), and the
possible adaptive advantage of this style of venation is
discussed by Zodrow & Cleal (1993). This morphological
change is accompanied by a proliferation of the group of
species, especially in the Bolsovian.

The abundance of the N. obliqua group then undergoes a
dramatic collapse in the topmost Bolsovian and it eventually
becomes extinct in the upper Westphalian D. This collapse in
abundance coincides approximately with the appearance of
the second group of neuropterids at the base of the Westphal-
ian D. In the Franco-Belgian Basin, this consists of just one
species, namely N. ovata, but elsewhere in Europe there are
other, very similar species which come in at about the same
level (N. flexuosa, N. plicata and N. ervedosensis). This is
near the top of the Upper Carboniferous succession in this
basin and so provides no direct evidence of the diversity of
these neuropterids at higher levels. However, in other areas
such as South Wales (Cleal 1978) and NW Spain (Wagner et
al. 1983, Wagner & Alvarez-Vazquez 1991) it is clear that the
group continues to be abundant at least through the Westphal-
ian D and Cantabrian, and in some cases beyond.

There is no direct evidence from the Franco-Belgian Basin
of the phylogenetic origins of the N. ovata group. There is a
possible precursor in the Bolsovian of the Intra-Sudetic Basin
(N. praeovata), but this throws little light on potential ances-

C.J. CLEAL AND C.H. SHUTE

\—"DRIER INTERVAL” = “|
i N. ovata

; nN N. willieres

0 '

i ron N. ghayei

! y_ VW N. heterophylla

N_ obliqua group

WeD

Fig. 21 Detailed abundance variations of Neuropteris species,
plotted against stages (using abbreviations shown in Fig. 2). In the
graph of the Neuropteris obliqua group, black represents N.
obliqua, fine stippling N. parvifolia, diagonal hatching N.
semireticulata, and coarse stippling Reticulopteris muenstert.
Based on data from Laveine (1967), determined from the
Franco-Belgian Basin. The shaded expansion of the N. ovata
curve reflects its proliferation in other areas.

tors. Almost certainly, the group evolved in an extra-basinal
habitat, possibly from a N. heterophylla-like ancestor.

Laveineopteris (Fig. 22). The laveineopterids may b

divided into two main groups: those with larger pinnules (th
L. tenuifolia/loshii group) and those with smaller pinnule:
(the L. rarinervis group). The larger pinnuled-group firs
appears in the Langsettian with L. loshii, which reaches it
acme in the upper Langsettian. At about the Langsettian,
Duckmantian boundary, however, it undergoes a significan
decline in abundance, and is replaced by a number of specie
with more elongate pinnules (L. tenuifolia, L. hollandica, L
jongmansii, L. morinii). These species, especially L. tenut
folia, remain abundant and characteristic elements of the
Duckmantian and Bolsovian, but then towards the top of th
Bolsovian decline sharply to become extinct in the lowe
Westphalian D.

The L. tenuifolia group shows a reduction in abundance
about the Duckmantian-Bolsovian boundary, but is otherwis
an important and characteristic element found in thos
stages. However, towards the top of the Bolsovian it unde
goes a second and this time terminal decline, finally becomin
extinct in the basal Westphalian D.

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 37

r
— "DRIER INTERVAL” ——

L. rarinervis

L. morinii

L. jongmansil

hollandica

tenuifolia

loshii

WeD

H
1
'
= uU c oO
[e} 2 mo wv
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|
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(Fig. 22 Detailed abundance variations of Laveineopteris species,
| plotted against stages (using abbreviations shown in Fig. 2).
| Based on data from Laveine (1967), determined from the
Franco-Belgian Basin. The shaded expansion of the L. rarinervis
| curve reflects its proliferation in other areas.

| The L. rarinervis group of very small-pinnuled species
shows a somewhat different distributional pattern. In the
Franco-Belgian Basin it starts in the Bolsovian, having possi-
bly originated from the slightly older L. nicolausiana. It
proliferates during the Bolsovian. In the lower Westphalian
D it appears to decline in the Franco-Belgian Basin, but this
is symptomatic of it being at the top of the Upper Carbonifer-
Dus succession here; elsewhere in the paralic belt of
20alfields, it continues to be abundant through into the
‘Cantabrian.

Veuralethopteris (Fig. 23). It is well known that this form-
yenus is restricted to the Namurian and Langsettian, a point
vhich is borne out by the Franco-Belgian data. The only
ther point of possible significance is that, compared with
many of the other neuropteroid taxa whose extinctions are
sormally marked by a gradual decline in abundance, the
xtinctions of most of the neuralethopterids is characterized
ly a sudden proliferation followed by a sudden decline.

aripteris (Fig. 23). According to Laveine et al. (1989), the
jlant that bore paripterid fronds migrated from China to
<a in the early Namurian. Elsewhere in Europe, it first
jppears in the Kinderscoutian, while in the Franco-Belgian
‘asin its lowest occurrence seems to be in the Marsdenian.
‘he stratigraphically lowest species is P. gigantea, which

|
|

'—— "DRIER INTERVAL” - =H
'

P. linguaefolia

P pseudogigantea

yo Ven P. gigantea

N. rectinervis

N. schlehanii

N. jongmansil

N. larischir

Fig. 23. Detailed abundance variations of Neuralethopteris and
Paripteris species, plotted against stages (using abbreviations
shown in Fig. 2). Based on data from Laveine (1967), determined
from the Franco-Belgian Basin.

extends through the rest of the Namurian, and proliferates in
the Langsettian.

At about the start of the Duckmantian, P. gigantea starts to
show a progressive decline, and is replaced by a new set of
species (P. pseudogigantea, P. linguaefolia). These remained
important elements of the Duckmantian and Bolsovian equa-
torial floras, except for a brief and temporary decline near the
Duckmantian-Bolsovian boundary. Towards the top of the
Bolsovian, however, these paripterids start a more significant
reduction in abundance, and they eventually become extinct
just below the base of the Westphalian D.

Macroneuropteris. Only one species of this form-genus
occurs in the Franco-Belgian Basin (M. scheuchzeri), and so
it has not been shown on the charts. The lowest occurrence of
M. scheuchzeri) here is in the upper Duckmantian, although
elsewhere it has been documented from as low as the upper
Langsettian (Pennines Basin — Cleal 1979). It reaches an
acme in the upper Bolsovian and then appears to decline.
However, it should be noted that elsewhere it remains an
abundant species through to the Cantabrian.

Palaeoecological controls on species distributions

From the above analysis of species distributions, a clear
pattern has emerged. Most significantly, there are two major
stratigraphical levels where changes occur:

1. The Langsettian-Duckmantian boundary. This marks (a)
the extinction of Neuralethopteris, (b) the start of the decline
of the early group of Neuropteris species, (c) the transition
from Laveineopteris loshii to the more elongate-pinnuled
laveineopterids (L. tenuifolia group), and (d) the transition
from Paripteris gigantea to P. pseudogigantea and P. linguae-
olia.

a The Bolsovian — Westphalian D boundary. This marks
(a) the extinction of Paripteris, (b) the decline and eventual
extinction of Laveineopteris, (c) the decline and eventual
extinction of Reticulopteris and Neuropteris semireticulata,
and (d) the sudden appearance and proliferation of the
second group of Neuropteris species allied to N. ovata.

It is clearly tempting to search for a palaeoecological
explanation for these two ‘events’, and we believe that such
an explanation can be found in the results of the coal ball
analyses summarized by DiMichele et al. (1985). Their model
was based on a number of different lines of evidence from the
peat-accumulating habitat vegetation, including species com-
position and the extent of the peat deposits. It seemed to
show that through the Late Carboniferous edaphic conditions
in the swamps would vary, with some periods of time being
slightly drier than others. In the middle Westphalian, for
instance, they found that some of the arborescent lycophyte
genera declined (e.g. Lepidophloios, Diaphorodendron) and
there was a corresponding increase in the Mesoxylon/
Mitrospermum-type cordaites, which they interpreted as indi-
cating rather drier conditions. From the point of view of our
study this is significant, as this drier interval ranged from
about the start of the Duckmantian to the end of the
Bolsovian, which exactly fits with the neuropteroid distribu-
tional patterns that we have found. To make this clear, we
have plotted this ‘drier interval‘ on the distributional charts in
Figs 21-23.

If the correlation between the coal ball data and the
neuropteroid distributions can be accepted, it has a number
of significant results:

1. Neuropteris species, except for those that developed a
significantly flexuous to pseudoanastomosed venation, were
mainly restricted to the wetter interval.

2. The development of flexuous, pseudoanastomosed and
eventually reticulate veining in Neuropteris/Reticulopteris
occurred when there was a change to drier conditions. It
would seem to have been caused by a fundamental change of
the genotype as, when conditions reverted to being wetter in
the Westphalian D, Reticulopteris was unable to reverse the
change.

3. The earliest known laveineopterid (L. loshii) was com-
monest at the time of wetter conditions in the Langsettian.
This was replaced as the dominant member of the form-genus
by the more elongate pinnuled forms (L. tenuifolia, L.
jJongmansii, L. hollandica, L. morinii) when conditions
became drier, at about the Langsettian-Duckmantian bound-
ary. The change was gradual and some pockets of L. loshii
persisted through to the early Bolsovian (for instance, the
well-known Duckmantian flora of the Barnsley Seam of
Yorkshire, U.K.).

4. The reversion to wetter conditions in the Westphalian D
coincided with the rapid decline and eventual extinction of
the elongate pinnule forms of laveineopterid.

5. The small pinnule forms of Laveineopteris (L. rarinervis)
appear not to be constrained by the same environmental
factors as the rest of the species. They first appeared in the

C.J. CLEAL AND C.H. SHUTE

drier interval of the middle Westphalian, but seemed equally
at home in the wetter conditions of the Westphalian D.
Macroneuropteris would seem to have been similarly unaf-
fected by the environmental change in the early Westphalian
D.

6. Like the laveineopterids, there was just one paripterid
species in the first wet interval (P. gigantea). It appears to
have many features in common (although it is not exactly the
same species — Laveine, pers. comm., 1992) with the pari-
pterids found in the upper Visean of China, which are
thought to represent the ancestral stock of this form-genus
(Laveine et al. 1989, 1992). According to the Laveine et al.
model, paripterids spread out westwards from China during
the very late Visean and early Namurian, along the northern
coast of the Proto-Tethys Ocean. It is likely that these early
paripterids favoured the wetter habitats of the lower delta
plains. It would thus not be surprising that the earliest
paripterid in Europe (P. gigantea) would also favour wetter
habitats.

7. Again, like the laveineopterids, on the change to drier
conditions in the early Duckmantian, the early species (P. |
gigantea) declined rapidly and was replaced by P. pseudogi-
gantea and P. linguaefolia. Both of these later species may
have been adapted to the drier conditions of the middle
Westphalian and did not survive the return of wetter condi-
tions in the Westphalian D. This resulted in the extinction of
the whole form-genus, although the group as a whole per-
sisted through to the lower Stephanian in the form of its
reticulate-veined cousin Linopteris.
8. The upper Duckmantian and lower Bolsovian has numer-
ous marine bands, indicating a change to lower delta plain
conditions (Guion & Fielding 1988). This coincides with a |
temporary decline in abundance of both the laveineopterids
and paripterids, which then recovered in abundance when
middle delta plain conditions returned in the middle and
upper Bolsovian. The levees were almost certainly of lower)
topography in a lower delta plain setting, and thus repre-|
sented wetter conditions than the levees of the upper Lang-|
settian and lower Duckmantian. This seems to confirm that
these mid-Westphalian laveineopterids and paripterids were
more abundant in drier conditions.

9. The neuralethopterids appear to have been totally}
restricted to the wetter conditions prevalent in the Langset-
tian. Unlike the laveineopterids and paripterids, they seemed
unable to adapt to the change to drier conditions in the
Duckmantian and became extinct.

The correlation between these events, identifiable in the|
adpression record, and the changes in the coal-swamp petri-
factions is remarkable, but it is evident that they are not sharp
events. For instance, the start of drier conditions probably
ranged through the lower part of the Duckmantian, while the
return of wetter conditions gradually developed from the
topmost Bolsovian to the lower Westphalian D. This is
suggested by the moisture curve given for coal-swamps by
DiMichele et al. (1985, fig. 8.1), but the much better evidence
that we have from the adpression record demonstrates it far
more clearly.

DiMichele et al. (1985) argue that the ‘wetter’ and ‘drier
conditions in their model refer to the edaphic conditions,
which in turn were responses to variations in climate. How-|
ever, whether these climatic changes were in the swamp
forests themselves, or in the hinterlands that supplied the
river-waters is not clear. That the changes can be identified

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 39

over wide geographical areas in North America and Europe
suggests that climate may well have been a major factor.
However, the temporary decline of the laveineopterids and
paripterids in the upper Duckmantian and lower Bolsovian,
suggests that the topography of the levees may also have been
a controlling factor.

Species diversities in other areas

As already stated, it is impossible to do the same type of

detailed diversity analysis in the other areas as we have done

in the Franco-Belgian Basin. However, there are a few points
which can be made on the distributions in some of these other
places.

It is well known that in Saar-Lorraine, Laveineopteris
tenuifolia becomes prematurely extinct in the upper Bolso-
vian (e.g. Laveine 1989). This is normally interpreted as a
response to an environmental change in this basin, repre-
sented by a predominantly arenaceous interval known as the
Geisheck Formation. From what we have learnt in the

_Franco-Belgian Basin, it is tempting to suggest that the

Geisheck Formation represents rather wetter conditions to
| that represented in the underlying Sulzbach Formation, in
»which L. tenuifolia occurs commonly.

Macroneuropteris scheuchzeri also becomes prematurely
| extinct in the Geisheck Formation of the Saar-Lorraine
|(Laveine 1989). This might be regarded as unexpected, as
-macroneuropterids in the Franco-Belgian Basin seem rela-
tively tolerant of environmental change. However, Bertrand
| (1930) suggested that the Saar-Lorraine representative of this
form-genus might not be taxonomically identical to that seen
in the paralic basins, having somewhat smaller pinnules with
only one (rather than two) basal lobe. Although this view has
not been widely accepted in the literature, the differences in
response to environmental change may support Bertrand’s
‘original contention.

Over much of Europe, Neuropteris sensu stricto is rare in
the Duckmantian and Bolsovian. A significant exception is in
NW Spain, where N. resobae occurs abundantly in the
Duckmantian Curavacas Formation (Cleal 1981). This still
‘fits in with the general pattern, however, as the Curavacas
Formation is a unit of fluviatile deposits in an otherwise
Marine succession (Martinez Garcia et al. in Martinez Diaz
1983) and would thus presumably have wetter edaphic condi-
tions than present in the coalfields of the paralic belt.

Over much of Europe, Neuropteris sensu stricto undergoes
a significant decline in the lower Stephanian. This is in
agreement with the DiMichele et al. (1985) model, as they
claim that a second (and this time more significant) drier
nterval started in the Cantabrian or early Barruelian in the
oal-swamp habitats. In a few parts of Europe, however,
Veuropteris remains a significant component in the upper
itephanian, such as Gard and La Mure (two of the coalfields
if the Massif Central), NW Spain, N. Caucasus, Donets and
he Alps. This may indicate that these areas were environ-
nentally wetter compared with the other parts of Europe and
he paralic coalfields of North America.

In most of the other parts of Europe, Neurocallipteris is the
ominant neuropteroid form-genus in the drier interval of the
tephanian. At least some also extend up into the Autunian,
hich DiMichele et al. (1985) claim represents a return to
fetter conditions. However, it is far from clear that these
asal Permian beds are indeed wetter and, at least in Europe,
hot supported by the increasing presence of red-beds.

Table 4 Results of regression and correlation analyses of extinction
(L) rates against numbers of species present (N).

——_—————————————————————

No. of species Total No.
of same form-genus _ of species
Regression equation L=0-57N+0-22 L=0-13N+0-48
Correlation coefficient (r) 0-7807 0-5402
Level of confidence that
correlation is significant 99-99% 99-47%
Coefficient of determination 60-94% 29-18%

Species diversity and survival

We have so far indicated that at least some of the variation in
diversity within the neuropteroid fossil record can be corre-
lated with Palaeozoic climatic fluctuations, and with varia-
tions in the volume of suitable strata. However, it is to be
expected that other factors may have had a role. One in
particular, which our data is suitable to test, is the degree to
which extinction rates were controlled by competition.

This has been tested by a regression and correlation
analysis of the numbers of species present in each stage
against the number of species of each form-genus that
become extinct in that stage. Two separate analyses were
performed, one using the total number of species present as
the independent variable, and the other using the number of
species of the particular form-genus present. In this way it
was hoped to determine whether competition within a form-
genus was a more important factor in determining extinctions
than competition generally within the neuropteroid complex
as a whole.

The results are summarized in Table 4 and Fig. 24. The first
thing that is evident is that extinctions are significantly
correlated with both the number of species of the same
form-genus and the total number of species. However, the
level of significance is much higher in the analysis using the
number of species of the form-genus. Also, the coefficient of
determination (the proportion of the variance in extinction
rates due to variations in species numbers) is much greater;
nearly two-thirds of the variance in extinctions could be
accounted for by the number of species of the same form-
genus present, while less than a third is accounted for by the
total species numbers.

From this, we conclude that competition was an important
factor controlling extinction rates of these plants, and that it
was greater between species of the same form-genus than
within the neuropteroid complex as a whole. The fossils
represent plants that grew in a fairly narrow band of habitats
and so some level of competition would be expected between
most of the elements represented. However, in such a setting
it would seem reasonable to expect that competition would be
greatest between those species that were closest genetically.
In this light, it would seem that the form-genera outlined in
this paper truly reflect the genetic relationships between the
parent plants, and thus support the essential robustness of the
classification.

PALAEOPHYTOGEOGRAPHY

All of the records analysed in this paper originate from what

Extinctions

10) 1 2 3 4 5 6 7 8

No. of species of same form-genus

* Laveineopteris
= Neurocallipteris

4 Sphenoneuropteris

X Neuropteris

C.J. CLEAL AND C.H. SHUTE

Extinctions

|
Total no. of species |
}

+ Macroneuropteris * Neuralethopteris

* Paripteris

|
i
Fig. 24 Regression of extinction rates against species numbers (parameters L against N of Table 3); (a) regression against number of species |

of same form-genus; (b) regression against total number of neuropteroid species. |

Cleal & Thomas (in Cleal 1991) refer to as the Europe
Palaeoarea, one of the subdivisions of the Eurameria Palaeo-
kingdom. There have been suggestions that this phytochorion
can be further subdivided based on the plant fossil record
(e.g. Gothan 1954). To investigate this possibility, we have
examined our data using cluster analysis, to see if any
palaeophytogeographical structure can be discerned. Our
data is obviously not entirely suited to such an analysis, as it
only represents a small portion of the total fossil assemblages.
On the other hand, our data has the merit of having been
critically assessed, and is thus preferable to some of the other
recently published palaeogeographical analyses, based on
uncritical literature trawls.

The database

Initially, we attempted to look at the data as a whole, using
an algorithm that could account for empty data points. This
was so the analysis could take into account species being
sometimes absent from an area merely because there is no
strata of the appropriate age there, rather than there being
any fundamental phytogeographical reason. However, the
results were disappointing, revealing little structure that
could be related to the geographical distribution of the areas.
It seemed a strong possibility that the empty data points may
have significantly distorted the results.

To overcome this, the data was split into five, stratigraph-
ically separate blocks. This reduced the number of empty
data points to a much lower and acceptable level. It also
allowed us to see if there was any stratigraphical variation in
the geographical patterns. The starting-point was taken at the
Chokierian, as there were too few neuropteroid species at
lower levels to provide any meaningful results.

Chokierian — Yeadonian. ‘This corresponds to most of the
Namurian and includes records from 11 areas. Margarito-
pteris multivenosa and Neuropteris bohdanowiczii were
removed from the original data matrix. These species are
only known from the Alportian, and strata of this age are

absent in 5 out of the 11 areas. It was thought that might
seriously distort the results. This left 6 species, on which the)
clustering was based.

Langsettian. Originally 15 areas were clustered based on 18
species. However, the records for South Limburg were
omitted, in order that this analysis would be in conformity
with that for the next stratigraphical interval (see below). |
Duckmantian — Bolsovian. Originally 14 areas were clus:
tered based on 24 species. The initial result showed a major
discrepancy with the position of South Limburg, which
appeared to cluster at a low level with Turkey, South Spai

and the Alps, rather than with the other areas of the oa
belt, as would be expected. On examining the data matrix, i
seemed likely that this might be due to the inadequacy of thd
data from South Limburg, and so we decided to omit it from
the analysis (and in consequence from that of the Langset:

tian).

Westphalian D — Cantabrian. 15 Areas were clustered|
initially based on 14 species. The results were initially unsat|
isfactory, showing what seemed to be a strong ‘chaining
pattern, indicative of poor structure in the data. However, b
combining the records of Neuropteris plicata with N. ovata
and of Laveineopteris piesbergensis with L. rarinervis,
rather better structure became evident (the taxonomic ratio
nale for combining these species can be found in the system
atic section of this paper, although at this stage we ar
reluctant to make formal proposals of synonymy until th
type material is subjected to a more rigorous morphologic
investigation).

Barruelian — Autunian. This corresponds to most of th
Stephanian plus the basal Permian. The initial data s
consisted of 10 localities and 14 species. However, Saxon
and the Pyrenees were excluded, as they only contain recor
from the Autunian, and would thus distort the analysis. Als
the record of Macroneuropteris scheuchzeri from the Inte
Sudetic Basin, and of Neuropteris schaeferi from |

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 4]

Lorraine were excluded. There are doubts about the
reliability of the former record (see comments in systematics
section) and the latter is based only on a single small
fragment. The final analysis was thus run on 8 localities using
12 species.

Results

The dendrograms produced by the five analyses are shown in
Fig. 25. Up to the Westphalian D, a relatively simple pattern
can be seen. Many areas contain neuropteroid assemblages of
relatively low diversity, in the Namurian consisting of Neur-
alethopteris schlehanii and Paripteris gigantea, these being
supplemented by Neuropteris obliqua in the Langsettian. In
the Duckmantian and Bolsovian, N. schlehanii disappears
from these low diversity assemblages, and P. gigantea is
replaced by P. linguaefolia. Against this background of low
diversity assemblages, however, there are two assemblage-
groups that are of significantly higher-diversity and, perhaps
significantly, correlate with the areas of greatest coal produc-
tion. These are shaded on the dendrograms, and may be
summarized as follows.

1. The Paralic Belt assemblages. These include the most

| diverse and abundant assemblages of neuropteroids, and
consistently cluster together with Jaccard Coefficients of 45 or
more from the Namurian to the Bolsovian. In the Namurian
\it includes most assemblages of northern, central and eastern
Europe, although there is some suggestion that there is an
area of even greater diversity, particularly of neuraletho-
pterids, in France-Belgium, NW Germany, U. Silesia and N.
Caucasus. In the Westphalian, however, the group as a whole
is limited to the paralic-belt coalfields of northern Europe
‘(NE Germany and Lublin are not included in the Langset-
tian, but this may merely reflect the limited data available
from these areas).
2. The intra-montane basin assemblages. Assemblages from
'Saar-Lorraine and the Intra-Sudetic basins take on a distinc-
tive character in the upper Duckmantian and Bolsovian.
| While including some taxa also found in the paralic belt
assemblages, many important constituents of the latter are
‘missing (e.g. Paripteris pseudogigantea, the Neuropteris obli-
\qua group, N. heterophylla, and Laveineopteris rarinervis).

‘The assemblages from the Iberian Peninsula also have a
distinctive character, usually clustering quite separately from
the rest of the areas analysed. Examining the database in
detail shows that they are mainly of very low diversity but, at
least in the Duckmantian-Bolsovian, include some endemic
taxa (Laveineopteris guadiatensis, Neuropteris resobae). A
further investigation into the Namurian and lower Westpha-
lian neuropteroids of Iberia may well produce interesting
results.

In the Westphalian D the pattern breaks down at lower
stratigraphical levels. Most areas form a relatively amorphous
2roup, which includes much of the old paralic belt, together
with the Intra-Sudetic Basin, NW Spain, Turkey and N.
Caucasus. The chaining structure evident in this cluster in the
lendrogram suggests that there is some non-homogeneity
vithin the group of areas, but that no clear subgroups are
‘ecognizable (although, the distinctive SW UK assemblages
vith Neuropteris flexuosa and Macroneuropteris macrophylla
ire positioned at one end of the chain). This partial reduction
n palaeophytogeographical provincialism appears to corre-
fate with the withdrawal or reduction of marine influence

|
|

from most of Europe (e.g. there are no marine bands above
the middle Bolsovian in the paralic belt), and thus the
disappearance of the marked distinction between the paralic
and intra-montane basins. The only notable exceptions to this
pattern in the Westphalian D are Saar-Lorraine (it no longer
clusters with the Intra-Sudetic Basin) and the highly distinc-
tive Zwickau assemblages.

In the Barruelian to Autunian, the cluster of areas with
most diverse assemblages again seems to correlate with the
major coal-producing areas, in particular the Massif Central,
NW Spain and the Intra-Sudetic Basin. Saar-Lorraine seems
to maintain its distinctive character, while N. Caucasus has
clustered quite separately because of the presence of a
number of apparently endemic taxa (although it has to be
recognized that the palaeobotany of this area is far from well
documented).

In conclusion, the most diverse Namurian to Bolsovian
assemblages occur in the coal-bearing paralic belt of northern
Europe. The coherence of this group of areas breaks down in
the Westphalian D, possibly as a result of the disappearance
of marine influence in these areas. In the Stephanian, a
second cluster of high-diversity areas appears in the intra-
montane coalfields of central and southern Europe. Saar-
Lorraine (together for a time with the Intra-Sudetic Basin)
retains a distinct character from these high-diversity areas, as
does the short-lived Zwickau Coalfield. In general, therefore,
the distribution of the neuropteroid complex supports the
conclusions of Gothan (1954), that there is a clear-cut distinc-
tion between the plant fossil assemblages of the paralic and
intra-montane basins. It might be tempting to use the results
to justify a formal palaeophytogeographical subdivision of
the Europe Palaeoarea into palaeoprovinces. However, such
a move would be premature before other plant fossil groups
have been subjected to similar analyses.

Endemism of individual form-genera

While there is clearly significant variation in the geographical
distribution of individual species, the same is not, on the
whole, so for the form-genera. Particularly the commoner
form-genera (Neuralethopteris, Paripteris, Neuropteris,
Laveineopteris) appear to be fairly evenly distributed. The
only significant exception seems to be Sphenoneuropteris,
which, throughout its range, has only been found in intra-
montane basins. Neurocallipteris is also mainly restricted to
intra-montane basins, but this is almost certainly just a
function of it being primarily a Stephanian and Autunian
taxon, in which paralic basins had all but ceased to exist in
Europe.

Neuropteroids from outside Europe

This study has been exclusively on records from Europe, this
being where these fronds are best known. However, there are
records from other areas of the world, which we will discuss
briefly here.

North America. The Carboniferous of eastern and central
North America belongs to the Eurameria Palaeokingdom. It
is to be expected therefore that similar if not identical
neuropteroids would be found here as in Europe. The
problem is that, other than in the Maritime Provinces of
Canada (e.g. Bell 1938, Cleal & Zodrow 1989), the Carbonit-
erous adpressions of North America have been very little

42 C.J. CLEAL AND C.H. SHUTE

: (a)
2 © S
sh Sec 8
$s sis g
5 8 & 8 3
Ss 8 S88 100 90 80 70 60 50 40 30 20 10 0
By ta SS SS
Poesy Ee ero. Poe eS ee ee eee eeEeEEee—E———E—E_—E—E
x x Lublin ki f.
Chokierian -
< x Svoge
Yeadonian
x x Pyrenees
x x< bd NE Germany
wi se Donets
DORE CSS IDS ew U. Silesia
“ « <x <x x France-Belgium
My ee NW Germany
Bee me N. Caucasus
x NW Spain
Soles Turkey
Ge (b)
SS s oS
3 Sen = 8 28 § 8
Sa SSNRB. ARS S SSN EBS 100 90 80 70 CONN SON nn 40)) nnIZONED WO 9
Sess SS ee SES RES ESS —_— aE ee ee re eee
SPS 2S Sess BSReESNS SS
SOSA SS we gqgers & esa es 8
RVers acs te ore re SOY thts ace eS mS FSS
Langsettian
x x x Lublin &
x SR RS ES ce eS Be eS Pennines
POL ADSe Te PS PR ORD opt SW UK
x ba RK or ee ee Donets
pvatede pee TESS EES ES. ES HEE U. Silesia
MR hd St Eb x Me Be Se be Be be Be : NW Germany
x <x Kx KX OK x x <x * KX XK XK France-Belgium
Sembee: Intra-Sudetic
he fa 8S Alps
Rs Svoge
x x x Turkey
x <~ x N. Caucasus
* * x x x NE. Germany
* x x NW. Spain
ee a a S. Spain
5 §
BS ~ =
2 S is & Ss RS BS (c
2 5 8 = & ww S S S Ss 8 5 )
Ss § 8 8 N S So 1s S) SN Sse oN SoS ks
SS ese eS eS SEC YES ER ERTS EES
SSSSESTSESSSETSESTSSS ZS EER? 100 «9 80 70 60 50 40 (30 20 a)
VS SRUSSRS SSeS RP SNES Sse SS US Se ee eee Ee EE
jg SSeS eseyyse geese eg aes se Ss
¢ x x x x NE Germany :
ts ets : Duckmantian -
rd mm me eM mS x mE i a
wat Bolsovian
x x x Posies ili Tait Enc Peto Tet fea Pennines
he SI RS ES RR RS RS EY ES a es oe bs bs te ce os France-Belgium
Ba SS a EA reg er eS ES os) ea ts C : NW Germany
ett eta br re Los ote decnad 2 tad * Donets
DAR OCI “other. CUE OSs PRLS t9G D U. Silesia
x hobs te Ee ae Ms x SW UK
x x x ca org x x Intra-Sudetic §
<x x x ts x Saar-Lorvaine &
Seer: he Svoge
ie : x x 5 Alps
id har Turkey
x x x S. Spain
x >< ted NW Spain

Fig. 25 Cluster analyses using geographical distributions of neuropteroid species (see text for details of methods of analysis); (a) Chokierian
to Yeadonian; (b) Langsettian; (c) Duckmantian to Bolsovian.

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND

LOWER PERMIAN 43

3 2 Ss S
SJ S by iS
Buip Sonepat dane oe alee (d)
RRS Sees iS) Sees eNS2 8 e
3 § & SSS S FR Ss 8
Ss § ea Ss 8 8
Ss § 25883 § N 100 90 80 70 60 50 40 30 20 10 0
PVPues=sezeRe Vey
Nishi: Pennines Westphalian D -
x x x Turkey _
Cantabrian
i Me RS Donets
CS Ie Sy Tes NW Germany
x x x xX France-Belgium
tat a U. Silesia
x x Intra-Sudetic
x x N. Caucasus
x x x x K NW Spain
Pa ES ES OS i SW UK
x * Saar-Lorraine
x < Alps
x os a S. Portugal
= N. Portugal
x x Zwickau
St <
S 38 8 3
Seca! Gere suey (ies (e)
aS. § Ge Ss SS) RRS
SePsNsssryse
&§ 8 § sss § g § $3 100 90 80 70 60 50 40 30 20 10 0
a Foe See RRS TS es SS SS eS Ee ee ae eee eee,
RS) rs ears Re Ec aie Gh
x x x x Intra-Sudetic Barruelian -
palate! adi toes ipa nine NW Spain Autunian
x MOS RS se x Massif Central
* x x x x*K N. Portugal

Saar-Lorraine
S. Spain
Donets

x N. Caucasus

Pig. 25 cont

studied, at least in recent years. There are some exceptions,
such as Darrah’s (1969) monograph on the Mazon Creek
lant fossils, and some useful records by Gillespie et al.
1975), Gillespie & Pfefferkorn (1976), Gillespie & Crawford
1985) and Gillespie & Rheams (1985). These indeed suggest
close similarity to the European assemblages. However, on
heir own they are not really sufficient to allow a comprehen-
‘ive assessment of the North American records, which is why
hey were not incorporated into the analysis presented in the
resent paper (for a further review of the North American
ecords, see Pfefferkorn & Gillespie 1980).
| The western part of North America in the Carboniferous
\as been assigned to two separate phytochoria, which may be
eferred to as the Cordillera Palaeoarea of the mid-west
ates and the Oregon Palaeoarea of the Pacific coastal area
Pfefferkorn & Gillespie 1980, Cleal & Thomas in Cleal
991). No neuropteroids have been reported from the
regon Palaeoarea. The Cordillera Palaeoarea is very poorly
‘ocumented, with the sole exception of the plant fossils from

|
|

(d) Westphalian D to Cantabrian; (e) Barruelian to Autunian.

the Manning Canyon Shale (Tidwell 1967). Of Tidwell’s
records, the most significant is of Neuropteris cf. pocahontas
White, which is undoubtedly a neuralethopterid similar to N.
schlehanii. However, his record of ‘Neuropteris’ gigantea is
undoubtedly incorrect (at least one pinnule in the figured
specimen has a basiscopic lobe — it may in fact be an elongate
neuralethopterid) and his ‘Neuropteris’ ampelina Tidwell is a
Eusphenopteris.

Gondwana. There are no neuropteroids recorded from the
Carboniferous of the middle and high palaeolatitudes of
Gondwana (the so-called pre-Glossopteris and early Glosso-
pteris floras — reviewed by Wagner et al. 1985). However, the
palaeoequatorial parts of Gondwana, such as the Mérida
Andes of Venezuela, the Djerada Basin of Morocco and the
Sud-Oronais region of Algeria, yield typical Euramerian-type
assemblages. The published records include species of Neu-
ropteris, Laveineopteris, Macroneuropteris, Paripteris and
Neurocailipteris (Jongmans & Deleau 1951, Jongmans 1952b,

Pfefferkorn 1977, Migier 1982). However, these are either
unillustrated records, or just show small fragments, which are
difficult to assess; their generic affinities are probably correct,
but any further statement will have to await a more complete
documentation.

Cathaysia. Although in very similar palaeolatitudes to
Europe during the Carboniferous, only a few neuropteroids
are found in China. The most significant from an evolution-
ary point of view is Paripteris, which seems to have first
evolved in China in the late Visean (possibly Brigantian) and
only later migrated west to Europe in the Namurian (Laveine
et al. 1989, 1992). The Chinese specimens have traditionally
been referred to as Paripteris gigantea (e.g. Li et al. 1974,
Yang et al. in Wagner et al. 1983). However, recent work by
Zhang et al. (1992) and Laveine et al. (1992) has shown that,
although similar, the Chinese material is not conspecific with
that from Europe and it awaits a new name.

There is also some evidence that Neuropteris may occur in
China. There are numerous records from the Upper Carbon-
iferous (thought to be approximately equivalent to the
Stephanian in the Heerlen Classification) of North China of
Neuropteris ovata. However, their veining is denser and the
pinnules more broadly attached to the rachis than the typical
Westphalian D specimens of this species from Europe, and
Gothan & Sze (1933) referred them to a separate species, N.
pseudovata. Wagner (1963) went further, to suggest that
there is a close similarity between these Chinese fossils and
the species which is now referred to as Neurocallipteris
neuropteroides. This clearly raises a difficulty as to the status
of the Chinese fossils, as there is no published evidence of
their cuticles to prove whether they are neuropterid or
neurocallipterid. In view of their relative high stratigraphical
occurrence, these Chinese fossils are in clear need of revision.

Li et al. (1974) described some fragmentary specimens
from the Namurian of China as Lopinopteris intercalata Sze.
Laveine et al. (1987) have argued that they may be very
closely related to Neuropteris obliqua. However, there will
have to be a more complete documentation of the Chinese
material before its taxonomic position can be confirmed.

Angara. There have been a number of records of Neuropt-
eris from this palaeokingdom (e.g. Neuburg 1948, Gorelova
et al. 1973). Among the more completely known species are
‘N.” pulchra Neuburg and ‘N.’ izylensis (Chirkova) Neuburg.
Although only a few specimens of these species have been
documented in the literature, and the illustrations of these
are mostly poor, they demonstrate certain significant features
of frond architecture: they have ultimate pinnae terminated
by a pair of pinnules and intercalated pinnules on the
penultimate racheis. These are characteristic features of the
form-genus Paripteris, although the pinnule form and vena-
tion is rather different from any of the European or Chinese
species. Also of possible paripterid affinity is ‘N. “ dichotoma
Neuburg, although this observation is based on the similarity
of its pinnules and venation to the European species P.
gigantea; little of its frond architecture has been documented.

Two species with very large pinnules (up to 70 mm long)
have been described under the names ‘N.’ siberiana Zalessky
and ‘N.’ balachonskiensis Gorelova. One specimen of the
former, figured by Neuburg (1948: pl. 31, fig. 1), shows
pinnules apparently with two basal lobes or incipient pinnules
(again, the quality of the illustrations make their interpreta-
tion difficult). A comparison with Macroneuropteris is thus

hinted at, but far more material needs to be examined before
this could be confirmed.
A rather unusual-looking species has been described as ‘N.’
ignotus Gorelova in Gorelova et al. (1973). It has very
tapered, subfalcate pinnules, spaced widely along a very wide
rachis, and is quite different from anything that has been
previously assigned to the neuropteroid group. A comparison
with the once-pinnate peltasperm frond Compsopteris is
possible, although without more complete material, prefer-
ably including cuticles, this affinity would be difficult to
confirm.
Most of the other Angaran species that have been assigned
to Neuropteris (e.g. “N.’ tomiensis (Zalessky) Radchenko,
‘N.’ orientalis Radchenko) are all too small and poorly
illustrated to assess. As far as it is possible to make out, other |
than some possible paripterids, no good examples of neu- |
ropteroid fronds have been described from these floras.

C.J. CLEAL AND C.H. SHUTE
|
|
|
|

Kazakhstan. ‘The Carboniferous plant assemblages found
here are intermediate in composition between those typical of |
Eurameria and Angara (Meyen 1987). According to both
Vakhrameev ef al. (1978) and Cleal & Thomas in Cleal
(1991), about half of both species and form-genera in the
Middle Carboniferous (in the Russian chronostratigraphy,
equivalent approximately to the Namurian and Westphalian
of the Heerlen Classification) of Kazakhstan are also found in
Europe, and include some neuropteroids. |

The best documented records of Carboniferous plant fos-
sils from here are by Radchenko (1954, 1985) and Oshurkova |
(1967). Other than some large, isolated pinnules from the
Upper Carboniferous (in the Russian sense, i.e. approxi-
mately Stephanian), identified as the Angaran species “Neu-
ropteris’ dichotoma Neuburg (see above), most neuropteroid-
like material originates from the upper Visean and
Namurian. The latter are all characterized by relatively small,
vaulted, lateral pinnules with a weakly developed midve
and a distinctive, round apical pinnule. The lateral pinnules|
vary to an extent in shape, from round to oval to subrectang-
ular with a round apex, and have been assigned to various
species including Neuropteris antecedens Radchenko non
Stur, N. heterophylla Oshurkova non Brongniart, N.
pseudoheterophylla Radchenko, N. bulupalganensis Rad-
chenko non Zalessky and N. karagandensis Borsuk. How-|
ever, these morphological variants are frequently found
associated together, and they almost certainly belong to one
and the same species. Goganova et al. (1992) have recently
described some remarkably complete examples of this species
and found that it is fundamentally different from Neuropteris.
They propose that the correct name is Cardioneuropteris
asiatica (Radchenko) Goganova et al. Although the fronds
are bipartite, producing tripinnate primary rachis branches,
there are no intercalated elements between the secondary
pinnae. Also, in close association were numerous
Aulacotheca-like sporangial clusters, which in Europe ar
normally associated with the frond form-genus Alethopteris.
It is clear that Cardioneuropteris is fundamentally differen
from any of the neuropteroid form-genera found in Europe.

Mention should be made of specimens recorded b
Oshurkova (1967) from somewhat higher (probably West
phalian equivalent) strata under the name Neuropteris e

qua. Unfortunately, only one extremely small fragment wa
illustrated (Ibid.: pl. 15 fig. 8), which is totally inadequate fo
taxonomic assessment.

It seems that, other than the possible paripterid ‘N.

NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 45

dichotoma and the inadequately documented N. obliqua, no
unequivocal neuropteroid form-genera (at least in the Euro-
pean sense) have been recorded from Kazakhstan.

CONCLUDING REMARKS

We are minded at the end of our study to quote from the

preface to John Woodward’s (1729) pioneering palaeonto-

logical study; Now, that I have been for some time engaged in

Mineral Studyes, with no small Application, ‘tis a Pleasure to

me to find that it has not been wholly without Fruit. When we

first started out on our project we intended it purely as a

means of testing the robustness of the taxonomic scheme

proposed by Cleal et al. (1990). However, we have ended up
on a much longer journey into the realms of palaeogeogra-
phy, biostratigraphy, palaeoclimatology and population
dynamics. Trying to improve the taxonomy of a group of
organisms, whether living or extinct, has its own internal
_ logic, but we discovered that is has also provided an improved
tool for understanding the pattern of the temporal and spacial
distributions of the species. The distributions of the indi-
- vidual species were of course mostly already known, but the
_ more general patterns were obscured by the wholly artificial
» generic taxonomy traditionally employed. Grouping the spe-
- cies into what seem to be more natural form-genera provided
» a context for at last seeing more clearly these more general
patterns; we have been able to see the trees for the wood!

This demonstration of its geological utility of course also
| adds further support for the essential ‘naturalness’ of the
_Tevised taxonomic scheme. That a group of species responds
| in the same way to environmental pressures does not prove
| that they are closely related. However, if the species are also
' morphologically very similar at both the macroscopic (frond
architecture) and microscopic (cuticles) levels, there must
iclearly be a strong likelihood that they are a genetically
homogeneous group. There will always be the potential for
convergent evolution to confuse the issue, especially with
| organs such as leaves, but by using as many morphological
| characters as possible it should be possible to detect this. A
case in point is the close gross-morphological similarity
between the mainly Westphalian D to Barruelian Neuropteris
ovata and the mainly Stephanian C to Autunian Neurocalli-
|pteris neuropteroides. Some authors have gone as far as to
‘suggest that the latter is a descendant of the former (e.g.
Wagner 1963). However, their epidermal structures are very
different, as are their apparent responses to environmental
changes within the forests, and it is almost certain that the
‘similarity in gross morphology merely represents convergent
evolution.

Our study provides clear evidence of the long-known but
often forgotten fact, that there is a close symbiotic relation-
_|ship between the study of plant fossils and geology; the fossils
cannot be properly understood without an understanding of
the geological (sedimentological, stratigraphical, palaeogeo-
graphical) context in which they are found. Equally, the plant
fossils provide invaluable palaeoecological, biostratigraphical
and palaeophytogeographical data for improving our under-
standing of the geology. This information can then be
re-cycled back to improve our understanding of the original
vegetation (Cleal 1991: 223). As our study has demonstrated,
this iterative process is dependent on the availability of a

robust taxonomy, not only at the rank of species but also of
form-genus. Obviously, a form-genus cannot be the exact
equivalent of a whole-plant genus, being based only on a
single plant organ. Nevertheless, the aim should be to make a
form-genus as near as possible to a phylogenetically coherent
concept (Cleal 1986), and this can only be achieved by
detailed morphological and taxonomic study of the fossils.

ACKNOWLEDGEMENTS. We are deeply indebted to the following for
providing information on some of the species referred to in the study,
and with which we were not previously familiar: Professor J.-P.
Laveine (Université Science et Techniques, Lille), Professor R.H.
Wagner (Jardin Botanico, Cordoba), Professor M. Barthel (Museum
fiir Naturkunde, Berlin), Professor J.H.F. Kerp (Westfalische
Wilhelms-Universitat, Munster), Dr. H.W.J. van Amerom (Geolo-
gisch Bureau, Heerlen) and Dr. Z. Siminek (Ustiedni tstav geolog-
icky, Prague). Professor Barthel and Dr Simtinck, together with Dr.
A.C. Scott (Royal Holloway and Bedford New College, London),
are particularly thanked for providing some of the photographs
illustrated in this paper. The remaining photographs were produced
by the Photographic Unit of The Natural History Museum, to whom
we are grateful. We would like to thank Professor Barry Thomas
(National Museum of Wales) for permission to reproduce the recon-
struction shown in Figure 1. For assistance with statistical proce-
dures, we would like to thank Dr. A.B. Smith and Mr. C. Montcrieff
(The Natural History Museum, London), Professor M.C. Boulter
(N.E. London Polytechnic) and Dr. W.L. Kovach (University of
Aberystwyth). We gratefully recognize the contributions of Dr E.L.
Zodrow (University College of Cape Breton, Sydney), particularly in
the early phases, in the development of the taxonomy used in this
paper, and for useful discussions on this subject in general. Finally,
we would like to thank Dr. J. McEvoy (Newbury) for help with
computer facilities, and Mr. J.A. Cleal (Solihull) and Dr T.J. Ferrero
(The Natural History Museum) for producing the illustrated charts.

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a
®

| NEUROPTEROID FOLIAGE FROM CARBONIFEROUS AND LOWER PERMIAN 51

INDEX OF GENERA AND SPECIES
0 eee ee EEE a ee ee ee eee ee eee

This is an index of the systematic section, not the whole paper. Species which are regarded as ‘good’ in the sense used in this paper are shown in bold Roman
type, while earlier synonyms and combinations, and species based on inadequate type specimens, are in italics. The archaic spelling variant Nevropteris is not
distinguished in the index, and its entries are to be found under Neuropteris.

Alethopteris neuropteroides 24
' antecedens, Neuropteris 25
arberi, Neuropteris 31
. asturiana, Neuropteris 31
auriculata, Neurodontopteris 25, 32
Neuropteris 23, 25

beveridgei, Neuropteris 31
) blissii, Neuropteris 30
bohdanowiczii, Neuropteris 25
Sphenopteris 25
- bourozii, Neuropteris 30, 32
britannica, Macroneuropteris 23
Odontopteris 23
| brongniartii, Sphenoneuropteris 29
» bulupalganensis, Neuropteris 31

! chalardii, Neuropteris 20
_condrusiana, Neuropteris 25
cordata, Neuropteris 23, 25, 30, 31, 32
;

:delasti, Neuropteris 31
densifolia, Neuralethopteris 24
‘densinervosa, Neuropteris 25
)dimorpha, Neuropteris 29
Pseudopecopteris 29
Sphenoneuropteris 29, 30
‘dispar, Neuropteris 31
idoubravica, Neuralethopteris 24
Neuropteris 24

dufrenoyi, Neuropteris 25
)duprei, Neuropteris 30, 32
\dussartii, Neuropteris 30, 32

elegans, Sphenoneuropteris 29
\ervedosensis, Mixoneura 26
ervedosensis, Neuropteris 26

Filicites (Nevropteris) heterophyllus 26
| tenuifolius 22

Mabellinervis, Imparipteris 31

‘Texuosa, Neuropteris 26

‘ormosa, Neuropteris 20

gallica, Neurocallipteris 25
| Neuropteris 25
hermeri, Neuropteris 23, 27

Neuralethopteris 24
Neuropteris 20
jugosa, Neuropteris 31

kosmannii, Neuropteris 31

lanarkiana, Neuropteris 26
larischii, Neuralethopteris 24
Neuropteris 24
lata, Neuropteris 24
Laveineopteris guadiatensis 20
hollandica 20
jongmansii 20, 26
loshii 20, 22, 26, 30
morinii 22
nicolausiana 22
piesbergensis 22
rarinervis 22, 30
tenuifolia 22, 23, 26, 30
linguaefolia, Neuropteris 28
Paripteris 28
linguaenova, Neuropteris 28
Paripteris 28
longifolia, Neuropteris 24
loriformis, Neuropteris 24
loshii, Laveineopteris 20, 22, 26, 30
Neuropteris 20
lubnensis, Neuropteris 32

Macroneuropteris britannica 23

macrophylla 23

scheuchzeri 23

subauriculata 23
macrophylla, Macroneuropteris 23

Neuropteris 23
maltbyensis, Neuropteris 28
Margaritopteris multivenosa 23
marginenervis, Neuropteris 26
matheronii, Neuropteris 31
mathieui, Neuropteris 25
Mixoneura ervedosensis 26

grandifolia 31

muensterifolia 31

polyneura 27

praeovata 27

raymondii 30

wagneri 30

auriculata 23, 25
beveridgei 31
blissii 30
bohdanowiczii 25
bourozii 30, 32

bulupalganensis 31

chalardii 20
condrusiana 25

cordata 23, 25, 30, 31, 32

delasii 31
densinervosa 25
dimorpha 29
dispar 31
doubravica 24
dufrenoyi 25
duprei 30, 32
dussartii 30, 32
ervedosensis 26
flexuosa 20
formosa 20
gallica 25
germeri 23, 27
ghayei 26
gigantea 28
grangeri 26
guadiatensis 20
hemingwayi 20
heterophylla 26
hollandica 20
horrida 31
jongmansii 20
Jugosa 31
kosmannii 31
lanarkiana 26
larischii 24

lata 24
linguaefolia 28
linguaenova 28
longifolia 24
loriformis 24
loshii 20
lubnensis 32
macrophylla 23
maltbyensis 28
marginenervis 26
matheronii 31
mathieui 25

shayei, Neuropteris 26 montana, Neuropteris 32 montana 32
sigantea, Neuropteris 28 morinii, Laveineopteris 22 morinii 22

Osmunda 28 Neuropteris 22 multivenosa 23
Paripteris 28 muensterifolia, Mixoneura 31 nemejciana 29

var. 8, Osmunda 26 multivenosa, Margaritopteris 23 nicolausiana 22
sleichenites neuropteroides 25 Neuropteris 23 obliqua 22, 26, 27, 28, 31
/randifolia Mixoneura 31 obliqua forma impar 22, 26
rangeri Neuropteris 26 nemejciana, Neuropteris 29 ovata 23, 26, 27, 28, 30
juadiatensis, Laveineopteris 20 Sphenoneuropteris 29 ovata forma flexuosa 26
_ Neuropteris 20 Neuralethopteris densifolia 24 ovata var. grandeuryi 27

doubravica 24 var. pseudovata 27

emingwayi, Neuropteris 20

eterophylla, Neuropteris 26
eterophyllus, Filicites (Nevropteris) 26

ollandica, Laveineopteris 20
Neuropteris 20

orrida, Neuropteris 31

nparipteris flabellinervis 31
ovata 27
_ plesbergensis 22

‘ngmansii, Laveineopteris 20, 26

jongmansii 24

larischii 24

neuropteroides 24

rectinervis 24

schlehanii 24
Neurocallipteris gallica 25

neuropteroides 25, 27

planchardii 25
Neurodontopteris auriculata 25, 32
Neuropteris antecedens 25

arberi 31

asturiana 31

var. sarana 27
papilioniformis 25
parvifolia 27, 28
pilosa 27
planchardii 25
plicata 20, 27
praedentata 29
praeovata 27
pseudoblissii 30
pseudogigantea 28
pseudoimpar 31
pseudozamites 25

rarinervis 22

raymondit 30

rectinervis 24

rectinervis forma obtusa 24

resobae 23, 28

rytoniana 20

schaeferi 28

scheuchzeri 23

scheuchzeri forma minor 28

schlehanii 24

schlehanii forma rectinervis 24

schlehanioides 24

schiitzei 29

semireticulata 28, 31

squarrosaeformis 31

stipulata 27

subauriculata 23

subplicata 20

subsessilis 31

teberdensis 31, 32

tenuifolia 22

valdensis 27

venceslai 29

waltonii 31

willierei 28

zeilleri 31, 32
neuropteroides, Alethopteris 48

Gleichenites 25

Neuralethopteris 24

Neurocallipteris 25, 27
nicolausiana, Laveineopteris 22

Neuropteris 22

obliqua, Neuropteris 22, 26, 27, 28, 31
Pecopteris 26
forma impar, Neuropteris 22, 26
Odontopteris britannica 23
stradonicensis 30
Osmunda gigantea 28
gigantea var. 3 26
ovata, /mparipteris 27
Neuropteris 23, 26, 27, 28, 30

forma flexuosa, Neuropteris 26

var. grandeuryi, Neuropteris 27
var. pseudovata, Neuropteris 27

var. sarana, Neuropteris 27

papilioniformis, Neuropteris 25
Paripteris gigantea 28
linguaefolia 28
linguaenova 28
pseudogigantea 23, 28
schuetzei 29
veenti 28
parvifolia, Neuropteris 27, 28
Pecopteris obliqua 26
piesbergensis, /mparipteris 22
Laveineopteris 22
pilosa, Neuropteris 27
planchardii, Neurocallipteris 25
Neuropteris 25
plicata, Neuropteris 20, 27
polyneura, Mixoneura 27
praedentata, Neuropteris 29
Sphenoneuropteris 29
praeovata, Mixoneura 27
Neuropteris 27
pseudoblissii, Newropteris 30
pseudogigantea, Neuropteris 28
Paripteris 23, 28
pseudoimpar, Neuropteris 31
Pseudopecopteris dimorpha 29
pseudozamites, Neuropteris 25

rarinervis, Laveineopteris 22, 30
Neuropteris 22

raymondii, Mixoneura 30
Neuropteris 30

rectinervis, Neuralethopteris 24
Neuropteris 24
forma obtusa, Neuropteris 24

resobae, Neuropteris 23, 28

rytoniana, Neuropteris 20

C.J. CLEAL AND C.H. SHUTE

schaeferi, Neuropteris 28
scheuchzeri, Macroneuropteris 23

Neuropteris 23

forma minor, Neuropteris 28
schlehanii, Neuralethopteris 24

Neuropteris 24

forma rectinervis, Neuropteris 24
schlehanioides, Neuropteris 24
schuetzei, Paripteris 29
schiitzei, Neuropteris 29
semireticulata, Neuropteris 28, 31
Sphenoneuropteris brongniartii 29

dimorpha 29, 30

elegans 29

nemejciana 29

praedentata 29

wagneri 30
Sphenopteris bohdanowiczii 25
squarrosaeformis, Neuropteris 31
stipulata, Neuropteris 27
stradonicensis, Odontopteris 30
subauriculata, Macroneuropteris 23

Neuropteris 23
subplicata, Neuropteris 20
subsessilis, Neuropteris 31

teberdensis, Neuropteris 31, 32

tenuifolia, Laveineopteris 22, 23, 26, 30
Neuropteris 22

tenuifolius, Filicites 22

valdensis, Neuropteris 27
veenii, Paripteris 28
venceslai, Neuropteris 20

wagneri, Mixoneura 30

Sphenoneuropteris 30
waltonii, Neuropteris 31
willierei, Neuropteris 28

zeilleri, Neuropteris 31, 32

Bull. nat. Hist. Mus. Lond. (Geol.) 51(1): 53-72 Issued 29 June 1995

The Upper Cretaceous ammonite Pseudaspidoceras
Hyatt, 1903, in north-eastern Nigeria

P. M. P. ZABORSKI bk €1%¢¢60
Department of Geology and Mining, University of Jos, P.M.B. 2084, Jos, Nigeria
CONTENTS

| VaUHROYG LLL) er sepa be pSE RE SSBe SERUM cite Neen MCE BE ee PCa sccreassomicenmarconabeadasscee aceccnit en re 53
NikeeAshalkavandseinci gs aysecuions atssrcatertsesoeeer- 5 ee tea eee ee ee ns
SY SLE IMAC CESCLIPUONS mee seetr tree neretnrm mee atte oc Na ccs ahincienecre cee tea et eT eee, eh tears BESS
Pavnavily AN CANNITUSKOXCSIRVANTDIDYANE, GRORSOINS ~cooacbponnsbadecasbesasenéoonssocedecabepdpesbesueoarenaédercnbobGscccoeebeces 55
SublamilysEU ONMPE ALO GER AGWINAEN Cooper tc--ee ites saan eee ee eee SS
GonustESHUDASEID O CRRA SWclyattimenee names sere ics eee sae eee Eee eee EE ee 55
SEudas pid Ocerasipseudonodosorresl(@hoitat) mse steetee ee eee ee 57
BS EUMAS DIG OGErGSs OOLEAUI (StOMCZKa) meeereee eee eee eee ee ee 59
[ESL DILOCAES DURING REST soooancecooosets see se ocaoneneadedeone seaosod ocopposbopeeooeboaamncuasmnencusedeose: 63
LESAN AOCT ES LEDS RONAN cogecionasudcoooseeaeoee osc eeeesbeoorcn decd ioconobersascposodeenaseoeeeon sceshon aber 63
GenngBURROCERAS CobbanwHookséa Kennedy, -:es:5s5-ssss4.0 ce eee eee 67
UIA ROTEAERY GD), conn Sp oS eeEeECe OCG: PREC EERE DECOTH So neRERR RPE Peer RRRSRCER i cia eee ee om nei ne ioe 69
Sitiasi rap CALdS CUSSION aa a asset oer. sees ee oo oes eeGoe se cero sala dace ese meee es cate eeeROGeE eee 69
IRGIEINENESS | cenisoddangooodde sob eO aaa nee en eee CP CSgE CE ERE E EE Oe ENR ETE er Weert mune ie uli mares ee Ait flat ewe I ited 71

Synopsis. The following species of Pseudaspidoceras Hyatt occur in north-eastern Nigeria, from oldest to youngest:
P. pseudonodosoides (Choffat), P. footeanum (Stoliczka), P. paganum Reyment and P. flexuosum Powell. P.
pseudonodosoides is represented by paedomorphic populations. P. flexuosum marks the basal Turonian of the
region. The remaining species are all of Late Cenomanian age. P. pseudonodosoides occurs in beds correlatable with
the Neocardioceras juddii Zone in south-western New Mexico but the Nigerian sequence is expanded in comparison;
no equivalents of the horizons with P. footeanum and P.paganum are known in the former region. The Nigerian
material demonstrates the great potential of Pseudaspidoceras in detailed correlation over the Cenomanian-
Turonian boundary. Below the beds with Pseudaspidoceras, specimens probably referrable to Burroceras Cobban,

Hook & Kennedy occur.

INTRODUCTION

In recent years renewed attention has been paid to the
Cenomanian-Turonian ammonite faunas of north-eastern
Nigeria (Wozny & Kogbe 1983, Popoff et al. 1986, Meister
1989, Zaborski 1990, Courville et al. 1991, Courville, 1992).
Early descriptions were provided by Woods (1911), Reyment
1954a, 1954b, 1955) and, especially, Barber (1957, 1960),
dut these works generally lacked information concerning the
wecise stratigraphical occurrences of the material involved.
(he impetus for fresh studies has largely been provided by
he opening of the Ashaka Cement Company’s quarry some
00 km north of the town of Gombe. Here, unusually for the
egion, large ammonite collections can be made within a clear
tratigraphical context. Coupled with information gathered
rom scattered localities elsewhere in north-eastern Nigeria,
otably the Pindiga stream section, data from Ashaka have
nabled a regional ammonite biostratigraphy to be worked
jut (Zaborski 1990). North-eastern Nigeria is emerging as a

) The Natural History Museum, 1995

region of key importance in understanding the ammonite
succession across the Cenomanian-Turonian boundary. The
expanded sequences are dominated by vascoceratid faunas,
but also contain acanthoceratid ammonites of wider geo-
graphical distribution, allowing correlations to be made with
zonal schemes from north-western Europe and North
America. One of the genera having such value is Pseudaspi-
doceras Hyatt, 1903, which is represented by four species, in
ascending stratigraphical order: P. pseudonodosoides (Chof-
fat), P. footeanum (Stoliczka), P. paganum Reyment and P.
flexuosum Powell. In addition, forms probably referrable to
the closely related genus Burroceras Cobban, Hook &
Kennedy, 1989 occur lower in the sequence. The purpose of
this contribution is to document the Nigerian material and to
highlight its stratigraphical value.

The Ashaka and Pindiga sections

Although Pseudaspidoceras occurs widely in north-eastern
Nigeria, only at Ashaka and Pindiga have more than one

species been collected

in an observed stratigraphical

sequence. The locations of these sections and other localities
mentioned herein were shown by Zaborski (1990), while a
general account of the geology of the region was given by

Carter et al. (1963).

A continuous section made up of ammonite-bearing lime-
stones and interbedded barren shales of Late Cenomanian
and Early Turonian age is exposed at Ashaka (Fig. 1). The
Pindiga stream section is discontinuous and partly subject to
burial and re-exposure during the summer rains. Figure 1
shows those parts of the section that have been logged over

the period 1986 to 1992.

The Ashaka section is as follows (see Fig. 1; numbers in
parentheses are the equivalent horizons in Meister (1989):

Bed

Metres

Alternating shales and sandstones, the lat-
ter glauconitic, feldspathic and calcite-
cemented below, quartzose above and
with a 15 cm shelly, sandy limestone
towards the top

Glauconitic, gypsum-bearing clay

Pale grey, nodular limestone with clay and
gypsum laminae. Vascoceras harttit
(Hyatt), V. obscurum Barber, Pseudotis-
sotia nigeriensis (Woods), Eotissotia sim-
plex Barber

Blue-grey shales with gypsum

Yellow, nodular limestone with Thalassi-
noides burrows on upper _ surface.
Pseudotissotia nigeriensis, Eotissotia sim-
plex

Blue-grey shales with gypsum

Yellow, nodular limestone with gypsum.
Thalassinoides burrows on upper surface.
Pseudotissotia nigeriensis, Eotissotia sim-
plex, Wrightoceras munieri (Pervin-
quiere)

Blue-grey shales with gypsum

Greenish-grey nodular limestone, weather-
ing to creamy yellow, with clay and gyp-
sum stringers. Vascoceras sp.,
Pseudotissotia nigeriensis, Eotissotia sim-
plex

Blue-grey shales with gypsum

Glauconitic, calcareous clay with black
phosphate pebbles. Pseudaspidoceras
flexuosum Powell, Watinoceras aff. colo-
radoense (Henderson), Vascoceras pro-
prium proprium (Reyment), V. obscurum
Barber, Thomasites gongilensis (Woods),
Pseudotissotia nigeriensis, Wrightoceras
munieri and Choffaticeras sp.

Cream-brown limestone with Pseudotissotia
nigeriensis and Vascoceras proprium pro-
prium (30)

Blue-grey shales with gypsum .................

Cream-brown calcareous concretions,
forming a continuous layer in places
Dark grey to blue-grey shales
Glauconitic, calcareous clay

X (34)

W (34)

V (34)

U (32)

T2 (30)

T1 (30)

S (28)

12-00

P.M.P. ZABORSKI

Hard, pale grey limestone. Pseudaspi-
doceras paganum Reyment, Vascoceras
proprium globosum (Reyment), Thoma-
sites gongilensis, Pseudotissotia nigerien-
sis

Calcareous, glauconitic, shelly clay, with
shale partings forming a more distinct
shale unit up to 50 cm thick in some
places

Massive, hard, grey-green limestone. Vas-
coceras sp., Thomasites gongilensis

Dark grey shales with gypsum; a 14 cm
horizon crowded with small thin-shelled
bivalves, bone fragments and small phos-
phatic pebbles occurs 15 cm below the top
in some places

1p Clay, with gypsum and scattered white cal-

careous nodules
Dark grey shales with gypsum

O (21-22) Hard, grey-green, nodular limestone in top

8-15 cm, rubbly limestone with shale part-

ings below. Pseudaspidoceras footeanum

(Stoliczka), Vascoceras proprium costatum

(Reyment), V. proprium globosum, V.

bullatum (Schneegans), V. nigeriense Woods,

V. cauvini Chudeau, Thomasites gongilensis,

ammonite gen. et sp. nov.

R (26)

Q (24)

N (20)
M (19)

Hard, grey, massive crystalline limestone ..
Rubbly, impure limestone with shale part-
ings. Pseudaspidoceras pseudonodosoides
(Choffat), Vascoceras sp. nov. aff. gamai
Choffat, V. cauvini

Hard, pale cream-grey, massive limestone .
Rubbly, impure limestone with shale part-
ings. Pseudaspidoceras pseudonodosoides,
Vascoceras sp. nov. aff. gamai, V. cau-
vini

L (18)
K (17)

J (16)
places
1 (14-15) Hard, grey limestone, cross-bedded in
places
H (12-13) Hard, grey limestone, cross-bedded in
places. Vascoceras cauvini

G(1i1) Massive, pale to dark grey limestone, pass-
ing laterally into bioturbated grey-green
limestone. Vascoceras cauvini

Rubbly limestone with interbedded harder
limestones. Burroceras? sp., Vascoceras

cauvini

F (10)

E (10) Rubbly, grey-green to dark grey limestone
with clay and gypsum stringers. Vasco-
ceras cauvini in upper part

D Massive, grey-green, glauconitic, quartzose
limestone. Exogyra-rich. Nigericeras gad-
eni (Chudeau)

Brown-yellow, calcareous sandstone

Ferruginous sandstone

Grey-green, fine-grained, poorly consoli-
dated sandstone becoming calcareous in
its upper part

>wWoO

~ mel

THE UPPER CRETACEOUS AMMONITE PSEUDASPIDOCERAS HYATT, 1903, IN NORTH-EASTERN NIGERIA

The Pindiga section, or more precisely that part of it which
has been seen, is as follows, from top to bottom, see Fig. 1
(numbers in parentheses are the equivalent horizons in

Popoff et al. (1986)):

Bed Metres
Blue-grey shales with gypsum (base of Pin-
diga Formation shale member)
V (22) Earthy, yellow-brown, glauconitic, calcare-
ous clay, with gypsum and white calcare-
ous nodules. Pseudotissotia nigeriensis,
ISCISXOTEL DO QUE’ csccectoaccog CEE RO EERCCA ORDO 0-15
Blue-grey shales with gypsum ................. 1-00
U(21) Pale grey-green, poorly bedded, impure
limestone. Pseudotissotia nigeriensis,
JE DYUBCOME OTD DIAS cococucseadangodenorecdeoqee0e 0-70
T (21) Brown-grey, poorly bedded, calcareous
shale, with gypsum and shale laminae.
Pseudotissotia nigeriensis, Eotissotia sim-
[DUB o accee tr Gamearno dssosccdcococa eee eA eRe ena 0:27
Blue-grey shales with gypsum ................. 0-15
18 (21) Cream-grey, irregularly bedded marl.
Pseudotissotia nigeriensis, Eotissotia sim-
[DUBE cer eapepeheap atise4c0¢ oc 08e0 aac een Rees 0-38
‘R(17) Light grey, poorly bedded limestone with
shale partings. Pseudotissotia nigeriensis.
7 cm hard, pale grey, nodular limestone
| CAPPS see acdeeteee eee te triniss oo isteslucuies tea 0-40
KO) Pale yellow, nodular limestone ............... 0-06
| Blae-oney Shales! qeecseeeeeeetere ses -icseceise cose. 0-04
| Hard, fine-grained grey limestone ........... 0-08
P Porous grey limestone. Thomasites gong-
ilensis, Pseudotissotia nigeriensis ........... 0-61
ILENE ONS. TAO) scaoaadooondadedquaconnaoaeeees 0-12
Hard, fine-grained grey limestone ........... 0-08
| Blue oneysshales|mesecserreeeece aecacecaceee seen 2-10
10) Intensely hard, grey, orange-weathering,
shelly limestone. ?Pseudaspidoceras
paganum, Vascoceras proprium globo-
sum, Thomasites gongilensis, Pseudotisso-
IL GATLLRCVLCTIS Stren ene erie: <tcei- coca e sist 0-22
\N (7) Hard, grey, laminated limestone with
Thalassinoides burrows on upper surface.
Pseudaspidoceras pseudonodosoides, Vas-
coceras sp. nov. aff. gamai, V. cauvini .... 0-18
Gy psumbang! ~-ieace rescence cscereieece neers 0-02
Blue-grey shales with gypsum ................. 0-10
M (7) Decalcified white limestone with Pseudaspi-
doceras pseudonodosoides, Vascoceras sp.
nov. aff. gamai and ammonite gen. et sp.
nov. preserved in a white clay matrix
LEP tO) SUR so. cist SPREE EEE REE RORY sie seiciecleencaene nes 0-05
Blue-grey shales with gypsum ................. 0-35
Gypsum band PE cence cisco ese ens 0-02
(7) Marly limestone. Vascoceras sp. nov. aff.
ELTON seep chips dco sous oboe Go ae pecieeeeoce ie ORECe 0-08
Gypsumiband . aeeeeeeteteecssecestces scenes 0-02
Islue-oney Shales Meereeer ses see- -caeee seer =~ <= 0-90
<(5,6) Hard, grey, nodular limestone ................ 0-15
BIWEAY STENES aocoaccobsccotoconodsnoceddnosebee 0-10
6.6) Hard, grey, shelly limestone. Reworked
Vascoceras cauvini on upper surface ...... 0-26

55
I (4) Roughly bedded, hard, grey, nodular lime-
stone with numerous Hemiaster ............ 0-45
H (4) Intensely burrowed, pale grey marl with
numerous Hemiaster. Burroceras? sp.,
WGSCOCELASICAUVINI Meee teen eee eee eee 0-23
G (4) Roughly bedded, impure limestone. Vasco-
CONAS | CAUVINIiacaans oh eeiccastas aceasta. sae 0-17
Blue-creyishales tans. cesesccsecaeee reesei te 0-06
F (4) Bryozoan biostrome in lower part, passing
upwards into 75 cm thick Plicatula bios-
trome, with Plicatula becoming less com-
IMOMUPWAnGSMrecren ssscnedsccckcdaeneatececeualel 1-21
Blue orevsshalesweren. sec. nnseuansteeeatedcaan eis 1-00
E Massive, hard, pale grey limestone .......... 0-13
Blue-oreyashaleswren.sces. saceseeeemerce neces 0-50
D Shelly, marly limestone with 1 cm shale
Pantin ppmEthematcd dle weceeeeeeeeseeeee ee eeee 0-07
Blue=oreysshalesteern. .).c ccs. eos-eecemesdenoaees 0-30
C nay WERS INTAESHONE. — croscoegudeoopocecdosodusso0000E 0-06
Blue oreygshalesmeree scare ea-ee sce ee teeecnee eee 0-20
B AFOUL JWAMESIOMNS cocoaonecvossogoocaconsnennc0ged 0-10
A (2) Rough-bedded, grey limestone with phos-
phatic particles at the base. Numerous
Exogyra; Metengonoceras dumbli (Cra-
gin), Placenticeras (Karamaites) cumminsi
(Cragin), Nigericeras gadeni ................. 0-42

Unit A at Pindiga is the “Exogyra Limestone’ of Barber
(1957). Units F-I are his ‘Echinoid Limestone’ while units O
and P are his ‘Gombeoceras Limestones 1 and 2’.

SYSTEMATIC DESCRIPTIONS

Repositories. Unless otherwise stated all the material referred
to herein is in the Department of Palaeontology, The Natural
History Museum, London. Only these specimens are indi-
vidually identified though many additional examples of Pseu-
daspidoceras pseudonodosoides and P. flexuosum from
Ashaka have also been studied.

Superfamily ACANTHOCERATACEAE Grossouvre, 1894
Family ACANTHOCERATIDAE Grossouvre, 1894
Subfamily EUOMPHALOCERATINAE Cooper, 1978
Genus PSEUDASPIDOCERAS Hyatt, 1903
(=Ampakabites Collignon, 1965a)

TYPE SPECIES. Ammonites footeanus Stoliczka, 1864; by
original designation.

REMARKS. Proposed by Hyatt (1903: 106), the genus Pseu-
daspidoceras has subsequently been discussed by Pervin-
quiére (1907), Freund & Raab (1969), Matsumoto (in
Matsumoto, Kawashita, Fujishima & Miyauchi 1978), Wright
& Kennedy (1981), Kennedy ef al. (1987) and Cobban et al.
(1989). It includes evolute ammonites with square to rectan-
gular whorl sections. There are distant to rather dense,
rounded to sharp, and rectiradiate to curved ribs in the
middle whorls. Umbilical, inner and outer ventrolateral
tubercles are present. Intercalated ribs frequently occur.
Freund & Raab (1969: 13) considered a wide lateral lobe to
be characteristic of the suture in Pseudaspidoceras. Matsu-
moto (1978) pointed out that Mammites wingi Morrow, 1935

56 P.M.P. ZABORSKI |

gl. j
20m
|
ASHAKA PINDIGA
NIGERIENSIS ZONE
15
Q |
p |
|
gl |
gl
=
— |
— }
™~
DO
PROPRIUM ZONE (concealed)
set i a |
10 — —— |
aa N
M |
L
}
K
J :
|
CAUVINI ZONE ;
:
5 a |
a |
ee
“a |
ae
GADENI ZONE
BIMA = |
SANDSTONE |. a —— /
0 4 E SN A
See
YOLDE
g! — glauconitic clay FORMATION

|
Fig. 1 Stratigraphical sections in the Pindiga Formation exposed at Ashaka Quarry and in the Pindiga stream. |

THE UPPER CRETACEOUS AMMONITE PSEUDASPIDOCERAS HYATT, 1903, INNORTH-EASTERN NIGERIA 57

and M. dixeyi Reyment, 1955 share this feature but these
relatively involute and stout forms have since been included
by Cobban & Hook (1983a) in their new genus Morrowites.
Hyatt (1903) assigned Pseudaspidoceras to the subfamily
Mantelliceratinae. Most subsequent workers have included
the genus in the Mammitinae (see, for example, Pervinquiere
1907, Reyment 1955, Wright 1957, Barber 1957, Matsumoto
1978, Wright & Kennedy 1981). Descriptions of the inner
whorls of P. flexuosum by Kennedy et al. (1987) and of P.
pseudonodosoides by Cobban et al. (1989), however,
revealed multiplication of the outer ventrolateral tubercles
and the presence of constrictions, demonstrating a close
relationship with Euomphaloceras Spath, 1923. Pseudaspi-
doceras is therefore best referred to the Euomphaloceratinae
Cooper. The type species of Ampakabites Collignon,
Kamerunoceras (Ampakabites) auriculatum Collignon
(1965a: 29, pl. 388, fig. 1662; pl. 389, fig. 1664), was regarded
as a synonym of P. flexuosum by Kennedy et al. (1987).
Collignon (in Cobban & Scott 1972: 81) had, himself, earlier
indicated that Ampakabites was better treated as a subgenus
of Pseudaspidoceras rather than Kamerunoceras.
Pseudaspidoceras has a stratigraphical range from Upper
_ Cenomanian to Lower Turonian. It occurs in Texas, New
_ Mexico, Arizona, Colorado, Mexico, Brazil, Germany,
' southern England, Portugal, Tunisia, Egypt and the Middle
| East, Algeria, Angola, Niger, Nigeria, Madagascar, southern
’ India and (?)Japan.

' Pseudaspidoceras pseudonodosoides (Choffat, 1898)
| Figs 2-5, 8, 14

| 1898 Acanthoceras(?) pseudonodosoides Choffat: 65, pl. 16,

figs 5—8; pl. 22, figs 32, 33.

| 1925 Mammites pseudonodosoides (Choffat) Diener: 175.

1957 Pseudaspidoceras sp. Barber: 11, pl. 25, fig. 8.

1969 Pseudaspidoceras cf. P. pseudonodosoides (Choffat);

Freund & Raab: 14, pl. 1, figs 10, 11; text-figs 4j-k.

| 1989 Pseudaspidoceras pseudonodosoides (Choffat); Meis-

. ter: 6, pl. 2, fig. 1; text-fig. 2.

| 21989 Pseudaspidoceras sp. Luger & Groschke: 372, text-fig.

| 1989 Pseudaspidoceras pseudonodosoides (Choffat); Cob-

ban, Hook & Kennedy: 40, figs 41, 81-83 (with syn-

) onymy).

/ 1990 Pseudaspidoceras cf. pseudonodosoides
Zaborski: figs 22a, b.

|
(Choffat);

| MATERIAL AND OCCURRENCE. Twelve specimens, C.93333,
—€.93335, C.93353-4, C.93573-5S, C.93982 from the Pindiga

Table 1 Morphometric data for Pseudaspidoceras
| pseudonodosoides (Choffat, 1898).

D Wb Wh U
|
C.47620 92 36 (39) 34 (27) 36 (39)
C.91232 91 40(44) 36 (39.6) 29 (32)
G4 7 - 31 (35.6) 35 (40)
C.93335 65 34 (52) 25 (38.5) =
31 (53.5) 22 (38) 20 (34.5)

|C.93757 58
|

\Dimensions (in mm). D, diameter; Wb, whorl breadth; Wh, whorl
height; U, umbilical diameter. Figures in parentheses are
dimensions as a percentage of the total diameter.

|
|

Formation, unit M, Ashaka; C.91232 from the lower part of
the Dukul Formation at Dukul (see Zaborski 1990: fig. 28);
C.93594 from the Pindiga Formation, unit M, Pindiga;
C.93756-7, from the Pindiga Formation, Deba Habe. In
addition a single specimen from the Pindiga Formation,
Bularaba (C.47620, see Barber 1957: 11) has been studied.
The species is very abundant in unit M at Ashaka and also
occurs in some numbers in unit K there. It occurs as a rarity in
units M and N at Pindiga. The Deba Habe specimens come
from a 10 cm limestone occurring less than 1 m below the
level at which Vascoceras proprium costatum and V.
nigeriense appear. No precise stratigraphical data are avail-
able for the Bularaba specimen.

DESCRIPTION. Most individuals have a maximum adult diam-
eter of 90-100 mm though some reach 120 mm. Whorl
breadth is slightly to distinctly greater than whorl height.

Only in C.47620 and C.93757 (Figs 5, 8) is the ornament of
the septate whorls well displayed. At a diameter of 6 mm in
C.93757 there are six prominent lateral bullae in the last half
whorl. They pass over the ventrolateral shoulders and merge
with bullate outer ventrolateral tubercles. Isolated outer
ventrolateral tubercles may be intercalated. In the last pre-
served whorl, up to a diameter of 58 mm, there are 9
umbilical tubercles. At first they are rounded and give rise to
robust, rounded, rectiradiate ribs on the low flanks. The ribs
bear a rounded inner ventrolateral tubercle but weaken as
they pass over the venter where they may branch or be
slightly convex. There are weak outer ventrolateral tubercles
but they disappear at a diameter of 35 mm. Intercalated
ventral ribs also occur; they may be as pronounced as the
main ribs or take the form of fine, convex riblets. The latter
type are more persistent, being present up to diameters of at
least SO mm. At diameters in excess of 45 mm the inner
ventrolateral tubercles weaken, then disappear. The umbili-
cal tubercles persist but become highly bullate and give rise to
broad, rounded ribs on the inner flank region.

In C.47620 (Fig. 5) there are 13 broad, rounded, rectiradi-
ate ribs in the whorl up to a diameter of 40 mm. Nearly all
arise at prominent umbilical bullae. They weaken in the
mid-flank region before bearing prominent, rounded inner
ventrolateral tubercles. The latter structures have disap-
peared by a diameter of 45 mm. There are no outer ventrolat-
eral tubercles at this stage. Though the earlier part of the
venter in this restored specimen is concealed, Barber (1957:
11) mentioned the presence of such tubercles on the early
whorls. Umbilical bullae persist onto the adult body chamber
where they become weaker, elongate structures giving rise to
irregularly developed, usually weak, concave ribs on the
flanks. The body chamber has a rounded outline in contrast
to the more rectangular, depressed section of the septate
whorls.

Umbilical bullae also persist onto the body chamber in
C.91232 from Dukul (Fig. 4) but here there are no flank ribs.
The material from Ashaka consists almost entirely of poorly
preserved body chambers which are generally smooth and
with a rounded whorl section. C.93335 (Fig. 3) and C.93982
are unusual in retaining a rather rectangular whorl section
and ribbing on the body chamber. In C.93982 the ribs extend
from umbilical bullae across the flanks and ventrolateral
shoulders where they terminate. In C.93335, however,
umbilical bullae are weakly developed but ventrolateral
swellings which give rise to rib-like structures on the inner
part of the flank are present. C.93574 (Fig. 2) shows broad,

P.M.P. ZABORSKI

ae ee PS se a ee ee a = ==

nyng ‘uoneUO, [NyNG “q “ep “Sly “TX “SEEES'O “Gq “BE “Bly “1X “PLSE6'O “4 ‘RZ “BIA “eYRYSY “WW MUN ‘uoNeUOY vsIpuld *¢ ©

rectiradiate ribbing on the adoral part of the phragmocone
: but the body chamber is smooth. Certain specimens, for
example that figured by Meister (1989: pl. 2, fig. 1) and
- C.93573, retain rather spinose ventrolateral tubercles on the
_ body chamber.

__ Specimens seen in units M and N at Pindiga are all portions
of the early whorls. One pathological specimen (C.93594,
Fig. 14) has a diameter of some 50 mm and an ornament of
_ umbilical bullae and abnormally developed inner ventrolat-
» eral tubercles, only one of the pair being present and dis-
) placed towards the siphonal line.

| Sutures in the Nigerian material described above are
) relatively simple with rather short, uncomplicated elements.
| The lateral lobe is often unusually narrow for the genus.
| When it is broader it is subdivided by a short median element.

REMARKS. The lectotype of P. pseudonodosoides (selected
by Cobban et al. 1989: 40) is the specimen figured by Choffat
(1898: pl. 16, fig. 5). It is fully septate, has a diameter of
about 90 mm, and is ornamented with strong umbilical
‘ tubercles, broad, rectiradiate ribs weakening in the mid-flank
)region, strong inner ventrolateral tubercles and, up to a
_diameter of about 60 mm, weaker outer ventrolateral
k tubercles. The whorls are distinctly broader than high and the
‘ venter is flattened to slightly concave. Choffat had only a few
“specimens at his disposal but recently Cobban et al. (1989)
' described a large collection from New Mexico which provides
-a wealth of information, especially regarding intraspecific
\ variation. These forms have whorls slightly to distinctly
) broader than high, the flanks being flat and the venters
| flattened to broadly rounded. Ribs are generally rounded but
may be sharp and narrow. There are 3—7 umbilical tubercles,
‘4-12 inner ventrolateral tubercles and 7—12 outer ventrolat-
‘eral tubercles in each half whorl. Ribbing is generally best
developed on the inner septate whorls while the outer ventro-
lateral tubercles weaken and disappear at diameters of 60-70
‘mm. Where they persist they may be expanded into oblique,
rib-like structures. There is a great range of adult sizes but no
Vevidence of size dimorphism; the diameter at the base of the
‘body chamber varies from 61-182 mm, while overall maxi-
/mum sizes are up to 300 mm.

The Nigerian collection conforms well with the lectotype,
vand, in Ornament and general shell proportions, with the
)more coarsely decorated material from New Mexico.
Although some of the latter specimens may have a compa-
table adult diameter, the material from north-eastern Nige-
Tia, without regard to its exact locality, has a consistently
smaller adult size of 90-120 mm. Material from Israel,
‘referred to P. cf. P. pseudonodosoides by Freund & Raab
(1969: 14-15), also includes forms reaching a diameter of
over 300 mm, though some show whorls higher than broad
and persistent outer ventrolateral tubercles and may be better
included elsewhere. Meister (1989: 9) suggested that P.
paganum (see below), a stratigraphically higher species, was
hypermorphic in comparison to P. pseudonodosoides. More
precisely, however, the north-eastern Nigerian poulations of
the latter species are probably paedomorphic. There is no
evidence of size dimorphism in this material, a similar adult
size is found in all the individuals from Ashaka, Dukul and
Bularaba. In this regard it is of interest to note the association
of P. pseudonodosoides with large numbers of Vascoceras sp.
nov. aff. gamai (=Vascoceras sp. juv. of Barber 1957: 27, pl.
5, figs 2, 4, 7; pl. 27, figs 10-15; Plesiovascoceras aff. gr.
Yhomi (Reeside) of Meister 1989: 11, pl. 4, figs 2, 3, 5;

THE UPPER CRETACEOUS AMMONITE PSEUDASPIDOCERAS HYATT, 1903, IN NORTH-EASTERN NIGERIA 59

Paravascoceras gr. evolutum (Schneegans) of Meister 1989:
14, pl. 5, fig. 4; text-fig. 10) at Ashaka, Pindiga and Deba
Habe. This Vascoceras has an adult body chamber homeo-
morphic with and of comparable size (about 100 mm) to that
in most of the co-occurring P. pseudonodosoides. The flank
ribbing in the early whorls of the former may also resemble
that in P. pseudonodosoides. The two are difficult to distin-
guish on the basis of poorly preserved material. The simpli-
fied suture in these P. pseudonodosoides even sometimes
approaches that in Vascoceras. Size and form of P. pseudono-
dosoides in north-eastern Nigeria may have been under
strong environmental control.

Pseudaspidoceras tassaraense Meister et al. (1992: 67, pl. 9,
figs 2, 4, 7; pl. 10, figs 1, 2; text-fig. 12) from Niger is also
adult at a diameter of only some 100 mm and loses its
ornamentation early to develop a rounded whorl section.
Meister et al. (1992) included the Pseudaspidoceras sp. of
Barber (1957: 11, pl. 25, fig. 8), here considered as P.
pseudonodosoides, in synonymy. P. tassaraense is closely
similar to the present material and is probably conspecific.

P. tassaraense is known only from the Monts I[guellala
region. Its precise stratigraphical level is difficult to deter-
mine in respect of other faunas in Niger. It occurs above
Nigericeras gadeni and Cibolaites? africaensis Meister et al.
(1992), the latter being unknown in Nigeria. Meister ef al.
(1992) inferred a position equivalent to a level between units
R and T at Ashaka for P. tassaraense, that is, well above the
horizons with P. pseudonodosoides there (units K and M).
Occurring alongside P. tassaraense in Niger are Nigericeras
Jacqueti involutum Meister et al. (1992: 68, pl. 4, figs 3-5;
text-fig. 14) and Vascoceras aff. gr. silvanense Choffat (Meis-
ter et al. 1992: 78, pl. 8, fig. 6; text-fig. 18). The V. cauvini
which accompany P. pseudonodosoides in Nigeria may
resemble N. jacqueti involutum in degree of compression and
lack of juvenile ornament but are consistently more evolute.
The inner whorls of certain Vascoceras occurring at the same
levels, however, are similar to the V. aff. gr. silvanense of
Meister et al. (1992) (see Zaborski 1991; fig. 18).

If P. tassaraense is conspecific with the present material
then its stratigraphical level is almost certainly lower than
that suggested by Meister et al. (1992). It would occur
between their Gadeni and Cauvini zones, that is equivalent to
their Pseudonodosoides to Evolutum zones in north-eastern
Nigeria (Meister et al. 1992: figs 22-26).

Pseudaspidoceras footeanum (Stoliczka, 1864)
Figs 6, 7, 9, 10

1864 Ammonites footeanus Stoliczka: 101, pl. 52, figs 1, 2.

1887 Ammonites pedroanus White: 217, pl. 22, figs 1, 2.

1915 Mammites (Pseudaspidoceras) footeanus(?) (Stolic-
zka); Greco: 208, pl. 17, fig. 5.

1936 Pseudaspidoceras pedroanum (White); Maury: 231, pl.
21, figs 1, 2.

1972 Pseudaspidoceras pedroanum (White); Reyment &
Tait: pl. 3, fig. 12.

1978 Pseudaspidoceras aff. pedroanum (White); Chancellor,
Reyment & Tait: 91, figs 8-10.

1982 Pseudaspidoceras footeanum (Stoliczka); Chancellor:
92, figs 2A, 24, 25.

1985 Pseudaspidoceras footeanum (Stoliczka); Howarth: 98,
figs 30-33.

1987 Pseudaspidoceras footeanum (Stoliczka); Kennedy,
Wright & Hancock: 38, text-fig. 4.

60 P.M.P. ZABORSKI

Figs 6,7 Pseudaspidoceras footeanum (Stoliczka). Pindiga Formation, unit O, Ashaka. Fig. 6a, b, C.93577, x0.67. Fig. 7a, b, C.93576,
x0.75. Fig. 8a, b Pseudaspidoceras pseudonodosoides (Choffat). Pindiga Formation, Deba Habe. C.93757, x1.

ig89,10 Pseudaspidoceras footeanum (Stoliczka). Fig. 9, Pindiga Formation, unit O, Ashaka. C.93578, x0.67. Fig. 10a, b, Pindiga
Formation, collected loose from the top of unit O, Ashaka. C.93362, x1. Figs 11, 12 Pseudaspidoceras flexuosum Powell. Pindiga
Formation, unit T2, Ashaka. Fig. 11a, b, C.93560, x1. Fig. 12a—c, C.93567, x1.

1992 Pseudaspidoceras gr. pseudonodosoides (Choffat);
Courville: pl. 1, fig. 2; pl. 2, fig. 1; pl. 3, figs 1,2

MATERIAL AND OCCURRENCE. Seven specimens, C.93309,
C.93362, C.93576-8, C.93764, C.93927, all from the Pindiga
Formation, unit O, Ashaka, except C.93362 which was
collected loose from the top of this unit. The species occurs
throughout unit O both in the rubbly limestone below and the
hard, nodular capping horizon.

Table 2 Morphometric data for Pseudaspidoceras footeanum
(Stoliczka, 1864).

D Wb Wh U
C.93577 210 70(33) 67 (32) =
C.93309 170 ~=— 65 (38) 56 (33) =
C.93576 135 58 (43) 55 (41) -
C.93362 65 30 (46) 27-5 (42) 22 (34)

Dimensions (in mm). D, diameter; Wb, whorl breadth; Wh, whorl
height; U, umbilical diameter. Figures in parentheses are
dimensions as a percentage of the total diameter.

DESCRIPTION. An evolute form with whorls a little broader
than high. The flanks are flattened and the venter flattened to
broadly rounded on the phragmocone, but the adult body
chamber may assume an evenly rounded outline. The maxi-
mum diameter attained is about 250 mm.

The earliest growth stages have not been seen. The small-
est specimen available (C.93362, Fig. 10) has a diameter of 65
mm and consists of half of one of the middle septate whorls.
The flanks are flattened, the ventrolateral shoulders sloping
and the venter broadly rounded. There are three prominent
and four feeble umbilical bullae giving off wide to narrow ribs
which weaken in the mid-flank region before passing into
prominent, rounded inner ventrolateral tubercles which are
also of irregular strength. The ribs curve forwards and
bifurcate over the ventrolateral shoulders before terminating
in well developed conical to bullate outer ventrolateral
tubercles which are often obliquely directed forwards. There
may be one or two pairs of outer ventrolateral tubercles
intercalated with those of the main ribs.

C.93576 (Fig. 7) has a diameter of 140 mm and consists of
about half of one of the later septate whorls. There are eight
umbilical bullae which give rise to low, broad, rounded,
mostly radial ribs which so weaken as to virtually disappear in
the mid-flank region. One of the ribs bifurcates with the
adoral branch curved distinctly forwards. Most of the ribs
bear prominent, rounded inner ventrolateral tubercles. Occa-
sional intercalated inner ventrolateral tubercles are present.
The ribs cross the ventrolateral shoulders and terminate at
weaker outer ventrolateral tubercles situated close to the
siphonal line. The latter tubercles vary from spinose to
rounded to bullate in shape and are situated opposite to or
slightly adoral of the inner ventrolateral tubercles. They
become less spinose during growth.

C.93309 has a diameter of about 170 mm and consists of the
adoral part of the near-adult phragmocone and the base of
the body chamber. The whorl section is subrectangular, the
flanks and the venter being flattened. Umbilical bullae give
rise to broad, low, unevenly developed ribs, effaced in the
mid-flank region and strengthening again as they cross the
ventrolateral shoulders where they pass into massive,
rounded inner ventrolateral tubercles, again of uneven

P.M.P. ZABORSKI

strength. The ribs terminate at very broad, low, rounded to
bullate outer ventrolateral tubercles.

Two adult body chambers are available. C.93577 (Fig. 6)
has a diameter of some 210 mm. Ornament declines markedly
and the whorl section becomes broadly and evenly rounded in
this individual. Towards the aperture there are narrow, sharp
ventrolateral ribs of uneven strength and spacing which are
asymmetrically developed on opposite sides of the specimen.
There are also weak, rounded, fold-like ribs crossing the
venter and extending onto the outer flanks. C.93578 (Fig. 9)
has a similar diameter. The style of ribbing typical of the
septate whorls extends onto the adapical part of the body |
chamber. The inner ventrolateral tubercles are highly spinose
here. On the adoral part of the body chamber the ribs |
become weaker and very widely spaced while the umbilical |
tubercles become weak, bullate, unevenly developed struc-
tures which may be twisted backwards. Flank ribbing is
virtually absent here though prominent, bullate inner ventro-
lateral tubercles persist and pass over the ventrolateral shoul-
ders as narrow, rib-like structures. The sutures in this
material are rather florid. The lateral lobe is very wide and
subdivided by a broad but fairly low median element.

REMARKS. The lectotype of P. footeanum (selected by
Wright & Kennedy 1981: 82) is the specimen figured by
Stoliczka (1864: pl. 52, figs la—c) (see also Kennedy et al.
1987: text-fig. 4). It has a diameter of over 250 mm. The
whorl section is quadrate. The ribs are generally radial and of —
uneven development on the later whorls. They arise from |
umbilical bullae but some fade before reaching the prominent |
inner ventrolateral tubercles. Outer ventrolateral tubercles
are persistent but adorally tend to merge with the bullate |
inner ventrolateral tubercles to form a ventrolateral rib-like
structure. A similar ornament is seen on a number of
fragmentary body chambers from Angola described by
Howarth (1985: 98, figs 31-33). The ornament of the lecto-
type and the Angolan specimens agrees well with that in the
later whorls of the Nigerian material. The collection from
Angola also includes one specimen showing the ornament of
the middle whorls (C.81073, Howarth 1985, fig. 30). This
individual closely resembles C.93362 from Nigeria though the
latter has outer ventrolateral tubercles located a little further
from the siphonal line and often elongated obliquely for-
wards.

Ammonites pedroanus White (1887: 212, pl. 22, figs 1, 2) is
a synonym of P. footeanum (see also Chancellor 1982: 94;) —
Bengtson 1983: 16; Howarth 1985: 98). Chancellor (1982: 95)|
suggested that P. paganum Reyment was also conspecific, |
along with the Nigerian specimen figured by Woods (1911:|
283, pl. 23, figs 1, 2). P. paganum is here considered to be a)
distinct species while Woods’ material belongs in P. flee
osum (see below).

The Pseudaspidoceras cf. footeanum of Wright & Kennedy
(1981: 82, pl. 21, fig. 3) has whorls distinctly higher than
broad. Its umbilical and inner ventrolateral tubercles are
larger than those of the lectotype and the outer ventrolateral
tubercles are closer to the siphonal line. In its smooth venter
and prominent inner ventrolateral tubercles it is like P| :
flexuosum. The specimen is reported as coming from r

stratigraphical level relatively high in the Lower Turonian
(Mammites nodosoides Zone at Dover) whereas the Nigerian
material is from the Upper Cenomanian. The Pseudaspi-
doceras sp. aff. footeanum of Matsumoto (in Matsumoto et al.
1978: 17, pl. 5, fig. 1) from Japan is a fragment only

THE UPPER CRETACEOUS AMMONITE PSEUDASPIDOCERAS HY ATT, 1903, IN NORTH-EASTERN NIGERIA 63

doubtfully referrable even to the genus.

Pseudaspidoceras footei var. grecoi Collignon (1965b: 176,
pl. E, figs 1a, b; Collignon & Roman in Amard et al. 1981: pl.
5, figs la, b) is a highly evolute form with a subquadrate

| whorl section a little higher than broad. Narrow, radial ribs,
_ effaced in the mid-flank, arise from umbilical bullae and bear
| prominent inner ventrolateral tubercles. Intercalated ribs
arise upon the outer flanks. All the ribs cross the ventrolat-
_ eral shoulders but do not reach the siphonal line. In its whorl
section and ornament this form is rather closer to P. paganum
_ (see below) than to P. footeanum.

Pseudaspidoceras reesidei Benavides-Caceres (1956: 468,
pl. 54, figs 1-4; text-fig. 51) is a moderately depressed species
) with an ornament similar to that in P. footeanum. The former
_ is, however, less evolute and its lateral lobe is subdivided by a
_ very low median element; in these respects it more resembles
_Morrowites.

__ The Acanthoceras(?) cf. footeanus of Choffat (1898: 66, pl.
6, fig. 5) seems to be adult at a diameter of less than 90 mm.

There are strong, rectiradiate ribs on the phragmocone while
‘the mainly smooth body chamber has weak, irregularly
developed, closely spaced ribs. The suture is unknown. This
‘form resembles the adults of P. pseudonodosoides from
Nigeria described above.

*Pseudaspidoceras paganum Reyment, 1954a
Figs 15. 16522523

1954a Pseudaspidoceras paganum Reyment: 253, pl. 4, fig. 1;
| text-figs 3h, 4.

1955 Pseudaspidoceras curvicostatum Reyment: 55, pl. 11,
| fig. 1; pl. 12; text-fig. 24.

1989 Pseudaspidoceras paganum Reyment;
| (pars), pl. 1, fig. 1; text-fig. 3.

11990 Pseudaspidoceras cf. flexuosum Powell; Zaborski: fig.
. 23 (only).

1991 Pseudaspidoceras flexuosum Powell; Courville et al.:
1041.

Meister: 6

(MATERIAL AND OCCURRENCE. Ten specimens, C.91275,
(0.93331, C.93537, C.93918-9, C.93920, C.93924-6, from the
Pindiga Formation, unit R (upper surface), Ashaka; C.93923,
ind the holotype (C.47422), are specimens collected loose
‘tom the Pindiga Formation, Pindiga but their matrix strongly
juggests derivation from unit O there.

fable3 Morphometric data for Pseudaspidoceras paganum
Reyment, 1954a.

D Wb Wh U
|
6.93331 146 55 (38) 58 (40) 45 (31)
7.93924 143 48 (33.5) 50 (35) 53 (37)
“93025 140 49 (35) 52 (37) 50 (36)
8.93537 135 46 (34) 50 (37) 52 (38-5)
93919 115 44 (38) 48 (39) 42 (36-5)

/imensions (in mm). D, diameter; Wb, whorl breadth; Wh, whorl
zight; U, umbilical diameter. Figures in parentheses are
(mensions as a percentage of the total diameter.

/ESCRIPTION. The whorls are quadrate or, more usually, a
ttle higher than broad. Maximum whorl breadth is at the
vel of the umbilical tubercles. The flanks converge slightly
oon the flattened to broadly rounded venter. At diameters

of less than 7 mm the whorls are ovoid and smooth. By
diameters of 20-25 mm there are narrow, fairly sharp, radial
to slightly convex ribs. They mostly arise at feeble umbilical
bullae and bear weak inner ventrolateral tubercles. They
curve forwards over the ventrolateral shoulders and termi-
nate in pronounced outer ventrolateral tubercles which are
elongated obliquely forwards. By diameters of 40-45 mm the
outer ventrolateral tubercles are more rounded in shape,
there being 2 or 3 for each inner ventrolateral tubercle. At
diameters of 50-60 mm the inner and outer ventrolateral
tubercles are of equal strength. At larger diameters the ribs
become unevenly developed. They are radial, convex or
flexuous in shape but may be effaced in the mid-flank region.
Most arise at variably developed umbilical bullae and all bear
bulbous to spinose inner ventrolateral tubercles which
become the most pronounced ornamental feature. Other ribs
arise in the mid-flank region and there may be pairs of
additional intercalated inner ventrolateral tubercles. Situated
adoral of the inner ventrolateral are rounded to clavate outer
ventrolateral tubercles. There are usually 1-3 pairs of addi-
tional outer ventrolateral tubercles between successive pairs
of inner ventrolaterals. These tubercles are of variable
strength and are sometimes asymmetrically developed. The
inner and outer ventrolateral tubercles persist as discrete
structures to the largest diameters seen, of nearly 150 mm. In
the later growth stages there may be weak, fold-like struc-
tures upon the flanks between the main ribs.

REMARKS. Chancellor (1982: 95) suggested that P. paganum
was a synonym of P. footeanum. The two are similar but
whorl breadth generally exceeds whorl height in the latter
while the opposite condition prevails in P. paganum. The
inner and outer ventrolateral tubercles also persist as discrete
structures to larger diameters in P. paganum; in P. footeanum
these tubercles take the form of bullate swellings on ventro-
lateral rib-like structures in the later growth stages. P.
paganum has a more rounded venter and its sutural elements
are more elongate and finely subdivided.

Pseudaspidoceras curvicostatum Reyment (1955: 55, pl. 11,
fig. 1; pl. 12; text-fig. 24) is a synonym of P. paganum (see
also Chancellor 1982: 92). The holotype (C.54801) is from the
Abazi River at Ezillo in south-eastern Nigeria. The species
was distinguished mainly on the basis of its strongly curved
ribs. As described above, however, this condition is also
found in the later growth stages in P. paganum.

The Acanthoceras cf. footeanum (Stoliczka) of Eck (1914:
196, pl. 17, figs 1, 2) is represented by a poorly preserved
specimen with whorls higher than broad and marked inner
and outer ventrolateral tubercles up to a diameter of at least
50 mm. This form may be most closely related to P.
paganum. As mentioned above P. footei var. grecoi Colli-
gnon also shows similarities with P. paganum.

Specimens referred to P. paganum by Barber (1957: 9) are
better placed in P. flexuosum (see below). Meister (1989: 8)
reported P. paganum from units O and R at Ashaka. All
members of the genus found in unit O during the present
work, however, are best referred to P. footeanum. The P.
flexuosum of Courville et al. (1991) are P. paganum.

Pseudaspidoceras flexuosum Powell, 1963
Figs 11-13, 17, 18, 20, 21

1902 Mammites footeanus Stol. spec. Petraschek: 144, pl. 9,
fig. 1.

EE — ——* a — ai ———— CT, Sats OOO — _ —_—

‘TX “pOSE6'D “PSIpulg “W yun ‘uoneui08.; esipuld “GeJJOUD) Saprosopouopnasd spsaz0pidsppnasg q “ep “1A ’
“TX “E9SE6'D “BYVYSY “ZL Wun ‘uoHeuUIO, esIpulg "[[aMog Wnsonxapf spdaz0pidsypnasg q ‘eg “Big

P.M.P. ZABORSKI

-16a,b Pseudaspidoceras paganum Reyment. Pindiga Formation, unit R, Ashaka. C.93331, x0.67. Figs 17,18 Pseudaspidoceras
\flexuosum Powell. Fig. 17, a, b, Pindiga Formation, unit T2, Ashaka. C.93366, x1. Fig. 18a, b, collected loose from the Pindiga Formation,
Pindiga. C.91276, x1. Fig. 19 Burroceras? sp. Pindiga Formation, unit H, Pindiga. C.93369, x1.

66 P.M.P. ZABORSKI

Figs 20, 21 Pseudaspidoceras flexuosum Powell. Pindiga Formation, unit T2, Ashaka. Fig. 20a—c, C.93367, x1. Fig. 21a, b, C.93911, x1.
Fig. 22a, b Pseudaspidoceras paganum Reyment. Pindiga Formation, unit R, Ashaka. C.93919, x1.

THE UPPER CRETACEOUS AMMONITE PSEUDASPIDOCERAS HYATT, 1903, IN NORTH-EASTERN NIGERIA 67

1911 Mammites (Pseudaspidoceras) sp. Woods: 283, pl. 23,
figs 1, 2.

1920 Pseudaspidoceras aff. pedroanum (White); Bose: 209,
plese tigers pla lS satioel:

1957 Pseudaspidoceras paganum Reyment; Barber: 9, pl. 1,
figs 1, 2; pl. 25, figs 5—7.

1963 Pseudaspidoceras flexuosum Powell: 318, pl. 32, figs 1,
9, 10; text-figs 2a—c, f, g.

1965a Kamerunoceras (Ampakabites) auriculatum Collignon:
29, 31, pl. 388, fig. 1662; pl. 389, fig. 1664.

1972 Ampakabites collignoni Cobban & Scott: 81, pl. 29, figs
1-3; text-figs 39, 40.

1987 Pseudaspidoceras flexuosum Powell; Kennedy, Wright
& Hancock: 34, pl. 2, figs 1-4, 8-13, 16, 17; text-figs
3A-C, 5, 6C, D, 7A-C.

1989 Pseudaspidoceras barberi Meister: 8, pl. 1, fig. 2; pl. 2,
figs 2, 5; text-fig. 4.

1989 Pseudaspidoceras flexuosum Powell; Cobban, Hook &
Kennedy: 41, fig. 91L.

1990 Pseudaspidoceras cf. flexuosum Powell; Zaborski: fig.
24 (only).

'1990 Pseudaspidoceras flexuosum Powell; Amédro in

| Robaszynski et al.: 264, pl. 17, fig. 1; pl. 18, fig. 1.

1992 Pseudaspidoceras barberi Meister; Courville: pl. 2, fig.
De

“MATERIAL AND OCCURRENCE. Nineteen specimens,
C.93366-8, C.93560-7, C.93911-7, from the Pindiga Forma-
‘tion, unit T2, Ashaka; C.91276, collected loose from the
' Pindiga Formation, Pindiga.

‘Table 4 Morphometric data for Pseudaspidoceras flexuosum
Powell, 1963 (see also Barber 1957: 9).

| D Wb Wh U

| €.93367 112 42(37:5) 46(41) 35 (31)
C.93567 98 42 (43) 42 (43) 30 (31)
C.93368 91 38 (42) 43 (47) 26 (28-5)
C.93366 name 311(37) 40 (48) 25 (30)
C.93566 81 28(34-5) 38 (47) 25 (31)
C.91276 75 29 (39) 34 (45) &
C.93565 72 32(44) 34 (47) sf
'C.93564 66 28 (42) 30 (45) 20 (30)
1C.93562 50 20 (40) 22 (44) =

Dimensions (in mm). D, diameter; Wb, whorl breadth; Wh, whorl
height; U, umbilical diameter. Figures in parentheses are
dimensions as a percentage of the total diameter.

DESCRIPTION. Although whorl height is generally a little to
markedly greater than whorl breadth, exceptional specimens
show quadrate whorls. The latter tend to have rather flat-
tened venters but in most cases the flanks are flattened and
the venters broadly rounded in this material.

C.93560 (Fig. 11) shows the ornament at a diameter of less
‘han 30 mm. There are 10-11 ribs in the last whorl which arise
it umbilical bullae. They bear prominent, bulbous to clavate
nner ventrolateral tubercles at which they loop forwards over
he venter and branch into pairs. The adapical rib of each pair
years weak, nodate outer ventrolateral tubercles located
lose to the siphonal line; the adoral rib is a simple untuber-
ulated structure curved convexly forwards.

The ornament of the middle whorls is variable. The ribs
nay be fairly broad and radial in disposition, distinctly

curved, or flexuous. They may be effaced in the mid-flank
region and often branch. All the main ribs arise at umbilical
tubercles, which are often of variable strength. They may be
bullate or spinose and twisted backwards. Each rib bears a
prominent, bulbous to bullate inner ventrolateral tubercle,
these structures outnumbering the umbilical tubercles where
the ribs branch. In other cases, ribs fade before reaching the
ventrolateral shoulder and here the umbilical tubercles out-
number the inner ventrolaterals. Across the venter each rib
bears a pair of weak outer ventrolateral tubercles located
close to the siphonal line and a little adoral of the inner
ventrolaterals. There may be additional weak intercalated
ventral ribs curved convexly forwards and without tubercles,
or additional pairs of isolated outer ventrolateral tubercles
may occur.

The outer ventrolateral tubercles may already have disap-
peared at diameters of about 50 mm or they may persist until
diameters in excess of 100 mm. After they have faded the
venter is broadly arched and flanked by prominent inner
ventrolateral tubercles. Weak, convexly curved ventral ribs
may persist up to diameters as large as 120 mm. No adult
body chamber is available. The sutures are complex with
narrow, elongate, finely subdivided saddles. The lateral lobe
is broad and divided, often asymmetrically, by a narrow,
elongate median element.

REMARKS. Chancellor (1982: 95) suggested that the Nigerian
ammonite described by Woods (1911: 283, pl. 23, figs 1, 2)
(specimen B3240 in the Sedgwick Museum, Cambridge) was
referrable to P. paganum and that this species was a probable
synonym of P. footeanum. He doubted, however, that all the
material referred to P. paganum by Barber (1957: 9)
belonged in P. footeanum. Similarly, Kennedy et al. (1987:
68) thought that Barber’s material was in part P. flexuosum.
Here, all this Nigerian material is included in P. flexuosum,
along with P. barberi Meister (1989: 8).

The most detailed previous description of P. flexuosum was
based on a large collection from west Texas (Kennedy et al.
1987: 34). The early whorls in the Nigerian material are
entirely comparable with those in the Texan specimens. The
minor differences in the later growth stages of the Nigerian
forms, occasional large whorl breadth and relatively coarse
flank ribbing, can be ascribed to individual variation. Consid-
erable inconsistency is displayed in these features by the
Ashaka specimens though they all come from a single S—6 cm
calcareous clay horizon. The variation is sufficient to encom-
pass Woods’ and Barber’s material. P. paganum has consis-
tently broader whorls, more pronounced, spinose and
persistent outer ventrolateral tubercles located a little further
away from the siphonal line, and a less complex suture.
Courville (1992: 423424) reported Pseudaspidoceras barberi
(= P. flexuosum) from unit U (his level 32) at Ashaka. The
fauna described by him, however, is that of unit T2 (upper
part of his level 30).

Genus BURROCERAS Cobban, Hook & Kennedy, 1989

TYPE SPECIES. Burroceras clydense Cobban, Hook &

Kennedy, 1989; by original designation.

REMARKS. Burroceras was proposed by Cobban, Hook &
Kennedy (1989: 37) for material from New Mexico transi-
tional in form and age from Euomphaloceras to Pseudaspi-
doceras. It combines the shell form and suture pattern of P.

Fig. 23a, b Pseudaspidoceras paganum Reyment. Pindiga Formation, unit R, Ashaka. C.93918, x1.
Fig. 24a,b Burroceras? sp. Pindiga Formation, unit F, Ashaka. C.93572, x1.

P.M.P. ZABORSKI

THE UPPER CRETACEOUS AMMONITE PSEUDASPIDOCERAS HYATT, 1903, IN NORTH-EASTERN NIGERIA 69

pseudonodosoides with the tubercle distribution of E.
euomphalum (Sharpe), siphonal tubercles being present.
Three species were identified, all of Late Cenomanian age:
B. clydense Cobban, Hook & Kennedy (1989: 38, figs 38,
79D-G, N-T) with a prominent ornament including ventro-
lateral horns and marked siphonal clavae; B. irregulare
Cobban, Hook & Kennedy (1989: 38, figs 39, 80S—V) with an
ornament of highly irregular strength; and B. transitorium
Cobban, Hook & Kennedy (1989: 39, figs 40, 79A-C,
80D-R) distinguished chiefly by its weak siphonal ornament.

Burroceras? sp. Figs 19, 24

MATERIAL AND OCCURRENCE. Three specimens,
C.93369-70, from the Pindiga Formation, unit H, Pindiga;
C.93572, from the Pindiga Formation, unit F (upper part),
Ashaka.

DESCRIPTION. C.93370 is part of a body chamber with a
diameter of some 120 mm. Umbilical bullae give rise to
robust flank ribbing. There are strong, rather clavate inner
ventrolateral tubercles but the ventral area is poorly pre-
served making identification of any ornament difficult.
_C.93369 (Fig. 19) is a more complete specimen with a
| diameter of 125 mm and consisting of one septate whorl and
_ about half a whorl of body chamber. There are narrow, radial
ribs of moderate strength on the septate whorl which arise
‘from umbilical bullae and bear prominent inner ventrolateral
‘tubercles. Again the ventral area is poorly preserved.
_ C.93572 (Fig. 24) is part of a septate whorl with a diameter
of about 130 mm. Whorl breadth slightly exceeds whorl
height. The flanks are flattened and the venter broadly
rounded. There are distant umbilical bullae of moderate
strength giving rise to low, rounded ribs which weaken in the
‘mid and outer flank regions. They bear fairly strong, rounded
‘inner ventrolateral tubercles at which they bend forwards and
become weak, broad structures before terminating at smaller,
rounded outer ventrolateral tubercles. There are vague traces
of siphonal swellings but it is difficult to determine if they are
real or preservational features. The suture shows a broad,
‘low median element subdividing the lateral lobe.

REMARKS. This material, from horizons low in the Ashaka
and Pindiga sections, differs from the earliest Pseudaspi-
ldoceras, P. pseudonodosoides, in the retention of discrete
outer ventrolateral tubercles until larger diameters. Unfortu-
nately, positive identification of siphonal ornament is not
dossible in these specimens and consequently they can only
de tentatively referred to Burroceras. Of the three species
oroposed by Cobban et al. (1989) in New Mexico the present
naterial most closely resembles B. transitorium in ornamen-
al style and suture pattern. In particular the siphonal
ubercles, located on a siphonal ridge which disappears on the
idult body chamber, are very weak, often barely noticeable
in this species.

JTRATIGRAPHICAL DISCUSSION

everal biostratigraphical schemes based on ammonites have
een put forward for the Cenomanian-Turonian beds in
orth-eastern Nigeria (see Reyment 1954a, 1954b, Barber
97, Wozny & Kogbe 1983, Popoff et al. 1986, Meister 1989,
saborski 1990, including a review, Courville et al. 1991).

That proposed by Zaborski (1990) is maintained here.
Briefly, five biozones were recognized, all defined at their
bases by the appearances of the nominal species. They are,
from oldest to youngest: a Zone of Nigericeras gadeni; a Zone
of Vascoceras cauvini; a Zone of Vascoceras proprium,
defined at its base by the appearance of V. proprium cos-
tatum, two further subspecies, V. proprium globosum (basal
Proprium to basal Nigeriensis Zone) and V. proprium pro-
prium (lower Nigeriensis Zone), being recognizable in the
region; a Zone of Pseudotissotia nigeriensis; and a Zone of
Wrightoceras wallsi Reyment. The stratigraphical distribution
of Burroceras(?) and the various species of Pseudaspidoceras
within this framework is as follows: Burroceras(?), lower part
of the Cauvini Zone; P. pseudonodosoides, upper part of the
Cauvini Zone; P. footeanum, lower part of the Proprium
Zone; P. paganum, basal Nigeriensis Zone; and P. flexu-
osum, lower part of the Nigeriensis Zone (see Fig. 1).

The best dated occurrences of Pseudaspidoceras elsewhere
in the world are in New Mexico and west Texas (see Hook &
Cobban 1981, Cobban & Hook 1983a, 1983b, Cobban 1984,
Kennedy et al. 1987, Cobban et al. 1989). In these areas only
P. pseudonodosoides and P. flexuosum are known, but in
south-western New Mexico Burroceras is also present. The
following biozones have been recognized in the Upper Cen-
omanian and Lower Turonian there (Cobban et al. 1989):

Zone of Mammites nodosoides
(Schliiter)

Zone of Vascoceras birchbyi (Cobban
& Scott)

Zone of Pseudaspidoceras flexuosum
Powell

Lower Turonian

Upper Cenomanian Zone of Neocardioceras juddii (Barrois

& Guerne)

Zone of Burroceras clydense Cobban,
Hook & Kennedy

Zone of Sciponoceras gracile
(Schumard)

Zone of Metoicoceras mosbyense
Cobban

Zone of Calycoceras canitaurinum
(Haas)

Burroceras clydense is the earliest known member of its
genus. B. irregulare and B. transitorium, along with Pseu-
daspidoceras pseudonodosoides, are Juddii Zone forms. P.
flexuosum characterizes the lowest Turonian zone recogniz-
able. There is, however, an hiatus horizon at the top of the
Juddii Zone in south-western New Mexico and it is at this
level that P. pseudonodosoides occurs; latest Cenomanian
beds equivalent in age to the Zone of Nigericeras scotti
Cobban in south-east Colorado and north-east New Mexico
are missing.

The Gracile Zone in New Mexico can be correlated with
the Nigerian Gadeni Zone as both contain Metoicoceras
geslinianum (d’Orbigny) (see Cobban et al. 1989, Zaborski
1990). The Nigerian Cauvini Zone is, at least in part, the
equivalent of the Juddii Zone as developed in south-west
New Mexico. The Burroceras(?) in unit F at Ashaka and unit
H at Pindiga cannot be identified to species level but appears
closest to B. transitorium, a Juddii Zone form. Equivalents of
the Clydense Zone cannot therefore be positively identified
in north-east Nigeria at present. Burroceras(?) in Nigeria is
associated with early Vascoceras cauvini which may closely

resemble V. barcoicense Choffat exile Cobban, Hook &
Kennedy (1989: 47, figs 47, 87Q-S, 89M-GG), at least in the
middle whorls, but the latter ranges from the Clydense to the
Juddii Zone in New Mexico.

Units K and M at Ashaka and M and N at Pindiga, which
contain the known stratigraphical range of P. pseudono-
dosoides, can be correlated with the upper part of the Juddii
Zone in south-western New Mexico. Lying disconformably
above in the latter region are sandy beds containing the P.
flexuosum fauna. This species occurs in unit T2 at Ashaka. As
explained below, there are no equivalents in south-western
New Mexico of units N to T1 at Ashaka. At Pindiga the
corresponding part of the sequence is largely unexposed,
though units O and P comprise a portion of it. The missing
part of the sequence in south-western New Mexico corre-
sponds to that containing P. footeanum and P. paganum in
Nigeria. In unit T2 at Ashaka P. flexuosum is accompanied
by large numbers of Pseudotissotia nigeriensis and in addition
Vascoceras proprium proprium, V. obscurum, Thomasites
gongilensis, Choffaticeras sp., Watinoceras aff. coloradoense
and Wrightoceras munieri (=W. wallsi of Meister 1989: pl. 28,
fig. 2). This fauna is of closely similar age to that from west
Texas described by Kennedy ef al. (1987) and taken to
characterize the basal Turonian Zone of Pseudaspidoceras
flexuosum there. Apart from P. flexuosum, Thomasites, V.
proprium proprium and W. munieri are notable common
elements. The appearance of P. flexuosum in the west
Texas-New Mexico area was suggested as a marker for the
base of the Turonian stage by Kennedy (1984) (see also
Cobban 1984, Birkelund ef al. 1984: 12). Hancock (1984,
1991) further suggested that V. proprium proprium might
perform the same stratigraphical function but at Ashaka this
form occurs in unit Tl, immediately below the first occur-
rence of P. flexuosum. On the basis of the appearance of P.
flexuosum, the base of the Turonian coincides with the base
of unit T2 at Ashaka. The top of unit T1 is a discontinuity
surface, overlain by a thin layer of diagenetic gypsum, while
unit T2 which has a high content of glauconite and phosphatic
matter is condensed. Construction of a correlation line
between the Ashaka and Pindiga sections indicates that the
Cenomanian-Turonian boundary in the latter occurs about 56
m above the base of the lowest limestone unit (A, see Fig. 1).
This portion of the section is unexposed. A single specimen of
P. flexuosum (C.91276, Fig. 18) has been found at Pindiga
but its precise horizon is unknown.

Courville et al. (1991), it should be noted, placed the
Cenomanian-Turonian boundary within what is regarded
here as the Upper Cenomanian since specimens of P.
paganum from both northern and southern Nigeria were
misidentified as P. flexuosum. Their P. barberi are synony-
mous with P. flexuosum.

The Cenomanian-Turonian boundary in north-eastern
Nigeria occurs in the lower part of the Nigeriensis Zone. The
earliest forms here referred to Pseudotissotia nigeriensis
(from unit R at Ashaka and unit O at Pindiga), however, do
not develop the typically tricarinate venter until relatively
large diameters of 50-60 mm. These individuals are morpho-
logically intergradational with their prolific associate Thoma-
sites gongilensis. There is little doubt that P. nigeriensis was
derived from the latter species (see also Barber 1957, Meister
1989). A clear distinction between the two cannot be made at
these stratigraphical levels. Meister (1989) evidently referred
the entire assemblage from unit R at Ashaka to T. gongilen-
sis. Undoubted examples of P. nigeriensis do, nevertheless,

P.M.P. ZABORSKI

appear before Pseudaspidoceras flexuosum; they are abun- |
dant in unit Tl at Ashaka. It may also be noted here that the |
identification of the Zone of Wrightoceras wallsi at Ashaka
and Pindiga by Zaborski (1990) was based on previous
reports of the occurrence of that species (see Barber 1957,
Popoff et al. 1986). Its presence has not been confirmed in
this work and the zone is not, therefore, indicated in Fig. 1.
Identifications of W. wallsi from unit T at Ashaka by Meister | -
(1989) are here regarded as dubious; the material is, at least |
in part, W. muniert.

If unit T2 at Ashaka represents the basal Turonian then
Pseudaspidoceras footeanum and P. paganum are Late Cen-
omanian in age. P. footeanum was assigned to the Lower
Turonian in Angola by Howarth (1985). Bengtson (1983: 44)
also recorded the species from the Lower Turonian in Brazil
but, in addition, listed similar forms from the high Cenoma-
nian. Records of P. footeanum, however, reveal little accu-
rate dating against other ammonites. The Nigerian examples,
from unit O at Ashaka, come from a stratigraphical level |
higher than that of the Juddii Zone but lower than that of the
Flexuosum Zone in south-western New Mexico. Unit O
contains an ammonite assemblage dominated by species of
Vascoceras. Prominent within the fauna are multitubercu- |
lated forms with simplified sutures (Nigericeras and Para-|
mammites of Barber 1957, and Vascoceras costatum
(Reyment) and Paramammites subconciliatus (Choffat) of}.
Meister 1989). This group shows a morphological gradation
into smoother ammonites referrable to Vascoceras nigeriense
Woods (1911: 281, pl. 21, fig. 6; pl. 22, figs 2, 3). The more}
strongly ornamented examples frequently resemble,
Nigericeras scotti Cobban (1971: 18, pl. 9, figs 1-4; pl. 18, figs}
1-9; text-figs 15-19), the nominal species for the topmost
Cenomanian zone recognizable in south-west Colorado and
north-east New Mexico.

The Pseudaspidoceras cf. footeanum of Wright & Kennedy)
(1981: 82, pl. 21, fig. 3) is reported as coming from a much|
higher stratigraphical level (the Mammites nodosoides Zone
of southern England) but, as mentioned above, probably|
does not belong in P. footeanum.

The exact stratigraphical level of the holotype of P.
paganum at Pindiga is uncertain. It is preserved in a hard,
orange-weathering limestone matrix closely matching unit O
there. The same is true of specimen C.93923, collected loose
at Pindiga. This bed is their probable source, especially as it
contains an ammonite fauna otherwise identical to that of
unit R at Ashaka, in which P. paganum is known to occur:
Thomasites gongilensis at its acme, Vascoceras propriu
globosum and the earliest Pseudotissotia nigeriensis. Units O
and P are the ‘Gombeoceras Limestones 1 and 2’ of Barbe
(1957) which make up his Zone of Paravascoceras costatu
(Reyment). Barber (1957: table 3) listed a number of specie
additional to those mentioned above which were said t
characterize these horizons. These forms, however, are prob
ably derived from loose limestone blocks found scattered in
gullies and fields at a stratigraphical level between units N”
and O at Pindiga. These blocks yield elements of the fauna
characterizing the Proprium Zone elsewhere including Vasco
ceras nigeriense.

Pseudaspidoceras pseudonodosoides is well dated in Israe
where it is found in the Vascoceras cauvini Zone (Freund
Raab 1969), a correlative of the Juddii Zone in north-wes
Europe, and in the New World (Lewy et al. 1984). Thi
stratigraphical occurrence is in accord with that in north
eastern Nigeria. In Portugal the species occurs at the base 0

THE UPPER CRETACEOUS AMMONITE PSEUDASPIDOCERAS HYATT, 1903, IN NORTH-EASTERN NIGERIA 71

the equivalent of the Juddii Zone but is recorded as ranging
into the Lower Turonian (Berthou 1984, Berthou et al. 1985).
P. footeanum is indicated as occurring low in the Juddii Zone.
Similarly, in Israel Freund & Raab (1969) indicated both P.
footeanum and P. cf. P. paganum as coming from the Cauvini
Zone alongside P. pseudonodosoides. These species have a
clear stratigraphical separation in north-eastern Nigeria and
such records require confirmation.
Amédro (in Robaszynski et al. 1990: 264) listed the order
_ of stratigraphical occurrence of species of Pseudaspidoceras
as: P. pseudonodosoides (Juddii Zone), P. flexuosum (basal
Turonian) and P. footeanum (Nodosoides Zone). This inter-
_ pretation seems to have been influenced by the description of
| P. cf. footeanum from the Nodosoides Zone by Wright &
' Kennedy (1981). However, as mentioned above, this speci-
men is closer to P. flexuosum but, even so, appears to occur
anomalously high.
| The expanded sequences across the Cenomanian-Turonian
boundary in north-eastern Nigeria provide the best docu-
' mented details available to date concerning the stratigraphi-
' cal distribution of Pseudaspidoceras as a whole. Clearly the
» genus has great potential value in detailed correlation at these
levels. Its occurrence in north-eastern Nigeria can be summa-
| rized as follows: P. pseudonodosoides occurs high in the
-Cauvini Zone equivalent to the upper part of the Juddii Zone
elsewhere in the world and is thus of Late Cenomanian age;
P. footeanum occurs in the lower part of the Proprium Zone,
probably at a level equivalent to the highest Cenomanian
‘Scotti Zone in parts of Colorado and New Mexico; P.
| paganum occurs in the basal part of the Nigeriensis Zone just
below the Cenomanian-Turonian boundary; P. flexuosum
occurs in the lower part of the Nigeriensis Zone equivalent to
a level marking the basal Turonian in west Texas and New
‘Mexico. Successive species show an increasing whorl com-
pression, increasingly delicate ornament and an increasing
isutural complexity.

—— Field work carried out in connection with this
work was largely funded by research grants from the Universities of
‘Maiduguri and Jos, Nigeria. Drs M. K. Howarth and H. G. Owen
and Messrs D. Phillips and S. Baker provided help in many ways.
_ Photographs were provided by the Natural History Museum (Lon-
jJon) Photographic Unit.

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The pterodactyloids from the Purbeck
Limestone Formation of Dorset

STAFFORD C.B. HOWSE and ANDREW R. MILNER
Department of Biology, Birkbeck College, Malet Street, London WC1E 7HX

CONTENTS
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Wenvicalivertebract seeseast ithe eae et ecw eens aete sce SREB EO ciate CRM SRS bs Sasi eee: See eee eens aecee setae 80
Hamilvyi@inmithocheimd acts celevee wernt ester sees see eicie= ais esis ecto sats eae he ates Bee CLS em mere sae eee a 83
GenusiOrnihocheinus! Seeley waa eecercrec terres ts ger Fe aie es athe eaeRE SRD AR ELE te bate faerie adapta em eee ae sete saeecienaicn 83
GOIN OCICITUSIBSP AN iets <p ooo el cots eiaise oes Salsas ne na otae ee rate Be sare aaleigan spies aN moeeriasaapeperbbs rine saettlnentins 83
Rterodactyloideayin Geraci sedis mrs acteun <tsletiess Saas beeches = Seles vs sae aiepiais splaeieess'amas lapses ac eee oes ao sete eetam aac 85
Doratorhynchus validus (Owen) Seeley nomen Vanur 1.12... .....ceceeen eee ee nec eee teceeneenee een tenesseneensenees 85
Othempterodactyloidimatentalitrompbunbecka mare -eeeeseecessee Geet eeGeee tees chee seca cece ee eee eae ee seer ee ee 86
ND) IS CUSS 1) Fa i tats et aes eT ee eas ge eects See va ec nage se Wt ee vatican slr eee araere ateiatuciemten te sete 87
Chronologicalirangeror pierodactyloidyramulicsy eecssees-es see eeeeceeseeereceecrereee eens see etn ereees saree ace ease ecees 87
Mhewburbeck wertebrateiiaumal Aha. sceche attack tise = sce caeiooeetie cette selene verge cine nieces ceismmeeet Gop cebisaa cea Oateee semen 88
FRE LET CNCE Sirs ee Psat se eae eee aie oe ae Sees alee Roe sas sa aaenninnnssombanmeclecauseecea se smeeeesides seeaduansscaboatioseess 88

Synopsis. The pterodactyloid fauna of the Tithonian-Berriasian Purbeck Limestone Formation of Dorset is revised
taxonomically, using both previously described and new specimens. Three distinct taxa are present, namely two
ctenochasmatids, Gnathosaurus macrurus comb. nov. based on a mandible and Plataleorhynchus streptophorodon
gen. et sp. nov, based on a rostrum, together with an ornithocheirid ‘Ornithocheirus’ sp. A, based on a second
mandible. Other pterodactyloid material from Purbeck includes probable ctenochasmatid cervical vertebrae, and
indeterminate pterodactyloid teeth, a radius, a metacarpal, a tibia and several wing phalanges, the latter including
the type of Doratorhynchus validus which is a nomen vanum. The presence of an ornithocheirid in the Purbeck
Limestone extends the history of this type of pterodactyloid back to the Jurassic-Cretaceous boundary and

Issued 29 June 1995

Ben 21¢G4G/

Since the middle of the nineteenth century, the Purbeck
Limestone Formation has produced a rich continental verte-
orate assemblage. Determinate pterosaur material has rarely
deen collected however, and only three isolated elements
qave been described or figured in the literature. Unfortu-
lately, the taxonomic treatment of this material has not been
mtirely rational and it is necessary to review the systematic
ustory of the Purbeck pterodactyloids to explain the system-
itic conclusions followed here. In 1868, H.G. Seeley
»btained a pterodactyloid mandible and an elongate vertebra
tom a quarryman at a quarry near Langton Matravers,
Purbeck, Dorset. Seeley (1869) listed these two specimens
jriefly, assigning them the name Pterodactylus macrurus. At

) The Natural History Museum, 1995

INTRODUCTION

reinforces the view that the major diversification of this group took place in the Middle or Upper Jurassic.

this stage, Seeley believed the vertebra to be a caudal of a
long-tailed pterosaur, although he assigned it to Pterodacty-
lus, a short-tailed genus. Owen (1870) figured a phalanx from
‘near Swanage’ as the new species Pterodactylus validus, but
made no reference to it in the accompanying text. Seeley
(1875) described the mandible and vertebra under Owen’s
species name validus despite the circumstances that firstly, his
own macrurus was the senior specific name; secondly,
Owen’s figure had not been accompanied by a description;
and thirdly, there was no basis for associating such disparate
fragments. In his description of the vertebra, Seeley vacil-
lated between identifying it as a caudal and as a cervical.
Seeley also concluded that a new generic category was
required for morphological and geographical reasons, and
created the genus Doratorhynchus for these specimens. This
aggregation of the Purbeck material under the binomen

Doratorhynchus validus has been followed by most subse-
quent authors e.g. Woodward & Sherborn 1890, Seeley 1901,
Plieninger 1929, Wellnhofer 1978 and Howse 1986. However,
Delair (1958) had reseparated Owen’s and Seeley’s speci-
mens, following Lydekker (1888) in referring only the pha-
lanx to Ornithocheirus validus. Curiously however, Delair
retained the binomen Doratorhynchus validus for Seeley’s
two specimens, thus creating the insupportable situation of
two validus species existing in place of one.

By 1901, Seeley had confidently reidentified the elongate
vertebra as a cervical of a long-necked pterodactyloid, and
noted the resemblance of this material to Solnhofen forms
such as Cycnorhamphus, but no explicit comparisons were
made. Doratorhynchus validus has subsequently remained in
the literature as an enigmatic pterodactyloid, either placed
doubtfully with the ornithocheirids, probably on the basis of
the mandible (Wellnhofer 1978) or considered to be a pos-
sible early relative of the azhdarchids on the basis of the
cervical vertebra (Howse 1986, Wellnhofer 1991a).

Recently, Angela C. Milner and one of the authors (ARM)
recognised two undescribed pterodactyloid cranial fragments
in blocks of Purbeck Limestone, in the collections of The
Natural History Museum, London. The early history of the
two specimens is unknown but they are recorded as having
originally been part of the collection of the Corfe Castle
Museum, the museum of the Purbeck Society. This collection
passed to Dorset County Museum in 1894 when the Corfe
Castle Museum closed (P.C. Ensom pers. comm.). It
remained as a separate collection within the Dorset County
Museum collections until part of it was purchased by the
British Museum (Natural History) in 1958. The pterosaur
specimens were subsequently shelved among the miscella-
neous crocodilians, until recognised recently. Preparation
and study of these two specimens has permitted a reassess-
ment and systematic revision of the previously described
Purbeck pterodactyloid material. The opportunity is also
taken to record some of the other fragments of medium-large
pterodactyloids collected at Purbeck over the last century.
The material referred to in this work belongs to the following
institutions (acronyms in parenthesis): Department of Palae-
ontology, The Natural History Museum (formerly British
Museum (Natural History)), London (BMNH); Bayerische
Staatssammlung ftir Palaontologie und Historische Geologie,
Munich (BSPM); Dorset County Museum, Dorchester
(DORCM); Sedgwick Museum, University of Cambridge
(SMC).

STRATIGRAPHY

All the specimens reviewed in this work are from the Purbeck
Limestone Formation of Dorset. Most were collected in
quarries in the Swanage/Langton Matravers area, the excep-
tions being a fragmentary cervical vertebra, a metacarpal and
some phalangeal material from Durlston Bay. None of the
specimens is accompanied by precise data on the locality or
horizon of collection. All of the Durlston Bay specimens and
some from inland quarries are labelled as having been
collected from the ‘Middle Purbeck Beds’ which, if accurate
(and this is by no means certain), could mean an origin in
either the Upper Lulworth Beds or the Lower Durlston Beds.
The two specimens first described by Seeley lack even this

S.C.B. HOWSE and A.R. MILNER

horizon data and are merely recorded as deriving from the
Purbeck Limestone.

Until recently, the Tithonian-Berriasian boundary (and
hence the Jurassic-Cretaceous boundary) was taken as the
‘Cinder Bed’ (now Cinder Member), partway through the
‘Middle’ Purbeck Limestone Formation at the boundary of
the Lulworth Beds and Durlston Beds (Casey 1963, Rawsdlll
et al. 1978). However, Allen & Wimbledon (1991) have}
pointed out that current attempts to define the base of the
Berriasian with more precision may alter the horizon of this
boundary within the Purbeck Limestone Formation. Whereas
selection of the Subthurmannia subalpina Subzone as the)
base of the Berriasian would maintain the status quo, selec-|
tion of the Berriasella jacobi Subzone as the base of the)
Berriasian would shift the Tithonian-Berriasian an
close to the base of the Purbeck Limestone Formation. Thus.
at present, the pterodactyloid material discussed in this work
is ambiguously Late Tithonian or Early Berriasian and is)
treated as such here, but should the Berriasella jacobi Sub-
zone be ratified as the new base of the Berriasian, then the
Purbeck Limestone Formation and its fossil assemblages
would all be of Berriasian (basal Cretaceous) age. |

SYSTEMATIC PAtALON Oa PALAEONTOLOGY

Order PTEROSAURIA Kaup, 1834 |
Suborder PTERODACTYLOIDEA Plieninger, 1901
Family CTENOCHASMATIDAE Nopsca, 1928

DIAGNOsIS (After Wellnhofer 1978). Pterodactyloids wit
elongate, anteriorly rounded snout. Food-trapping dentitio
composed of a large number of long, pointed, outwardl
directed teeth inserted laterally in the jaw margins. T
tooth-row extends back for about half the skull length.

INCLUDED GENERA. Ctenochasma Meyer, 1851; Gnathosau
rus Meyer, 1834; Huanhepterus Dong, 1982; Plataleorhyn
chus gen. nov.

RANGE. Upper Jurassic — basal Cretaceous; Europe an
China.

Genus GNATHOSAURUS Meyer, 1834

TYPE SPECIES. Gnathosaurus subulatus Meyer, 1834 from th
Tithonian of Solnhofen, Bavaria, Germany.

|
DIAGNOsIS (After Wellnhofer 1978 with further points in
parenthesis). Relatively large ctenochasmatids with lon
slender skulls, jaw tips spatulate and anteriorly rounded
Premaxillae bear a low sagittal crest extending from one thir
of the skull length behind the snout tip to the level of th
orbit. Prominent dentition (30-32 teeth per ramus), extend
ing back to the level of the anterior edge of the naso
antorbital fenestra. The anterior spatula teeth are short an
anteriorly directed, the posterior spatula teeth being longer
From the 5th tooth back on each side, the teeth lengthe
substantially and are anteroposteriorly directed so that eac
overlaps outside the tooth in front. (The posterior teeth ar
much shorter). Teeth are arranged to give a food-grabbin
and ‘fish-basket’ dentition, with the teeth inserted on th
lateral edge of the jaw margin and the tips inwardly directed

}

|
|
|

THE PTERODACTYLOIDS FROM THE PURBECK LIMESTONE FORMATION OF DORSET

Naso-antorbital fenestra about twice as large as the orbit.

RANGE. Tithonian/Berriasian of Germany and England.

Gnathosaurus macrurus (Seeley) comb. nov. Figs 1, 2

1869
1875

Pterodactylus macrurus Seeley: 89-90 partim
Doratorhynchus validus (Owen); Seeley: 465-8 partim
(non Owen 1870)

1891 Doratorhynchus validum (Owen); Woods: 169 partim
(non Owen 1870)

1901 Doratorhynchus validus (Owen); Seeley: 173 partim
(non Owen 1870)

1929 Doratorhynchus validus (Owen); Plieninger: 27-28
partim (non Owen 1870)

1958 Doratorhynchus validus (Owen); Delair: 70-1 partim
(non Owen 1870)

1978 Doratorhynchus validus (Owen); Wellnhofer: 58
partim (non Owen 1870)

1986 Doratorhynchus validus (Owen); Howse: 318-20

partim (non Owen 1870)

LecroTyPE. SMC J5339, the anterior region of a mandible
of a medium-sized pterodactyloid, exposed in dorsal aspect
(Figs 1, 2). This specimen has not previously been figured.
Seeley (1869) based ‘Pterodactylus’ macrurus on two ele-
ments, this mandible and a cervical vertebra which thus
constituted the syntypes of this species. There is no reason to
associate these two specimens and the vertebra could belong
to either of the large ctenochasmatids described here.
Although only the vertebra has been figured previously, the
mandible is more critically diagnostic of a taxon, and we have
accordingly selected it as the lectotype of macrurus.

DIAGNOsIs. Species of Gnathosaurus in which the spatula on
the lower jaw bears 10 teeth (14 in G. subulatus) and in which
the main series of post-spatula mandibular teeth are nearly
laterally directed (antero-laterally directed in G. subulatus).

TYPE LOCALITY. Quarry near Langton Matravers, Purbeck,
Dorset (Seeley 1875). The precise locality is not known.

HORIZON. Purbeck Limestone Formation; Tithonian, Upper
Jurassic or Berriasian, Lower Cretaceous. The precise hori-
zon is not known.

DESCRIPTION. SMC J5339 comprises an incomplete man-
dible exposed in dorsal aspect (Figs 1, 2). The anterior region
is abraded, the middle region is well preserved and the
posterior ends of both rami are absent. The anterior regions
of the mandibular rami coalesce with a long narrow common
symphysis (Fig. 1). The length from the anterior tip of the
mandible to the broken end of the longest ramus is 308 mm,
the length of the symphysis is 122 mm, and the greatest width
between the diverging rami is 51 mm. The anterior extremity
of the symphysis is expanded into a small spatula about 22
mm wide and 23 mm long (Fig. 2A). This is rounded
anteriorly but then narrows more gradually posteriorly to

_ give an elongate shape. The spatula bears ten alveoli, five on

each side. Their arrangement indicates that the spatula teeth

_ were all directed anterolaterally. Behind the spatula, the

symphysis constricts to a width of 12 mm, then widens to 25
mm at the point where the rami diverge. At this point, the
Tami meet at an acute angle and completely ankylose, there
being no trace of a symphysial suture. The symphysial region
is relatively shallow and does not appear to have been

crushed post-mortem.

Vr apne
WEE Se

sym.t.

LT]

Fig. 1 Gnathosaurus macrurus (Seeley) comb. nov. SMC J5339,
lectotype. Anterior region of mandible in dorsal aspect.
Abbreviations: sp. = spatula; sym.t. = symphyseal trough. Scale
= 10 mm.

al.10

al.20

S.C.B. HOWSE and A.R. MILNER

4 temcal’s)
p
y a
ral 10

al.15

sym.t.

al.25

Fig. 2 Gnathosaurus macrurus (Seeley), comb. nov. SMC J5339, lectotype. (A) Symphyseal region as preserved. (B) Reconstruction of

anterior mandible in dorsal aspect. Abbreviations: al. = alveolus; d.f. = dental foramen; sp. = spatula; sym.t. = symphyseal trough. Scale

= 10 mm.

THE PTERODACTYLOIDS FROM THE PURBECK LIMESTONE FORMATION OF DORSET 77

On the dorsal side of the midline of the symphysis is a
symphysial trough, a groove about 2 mm wide and between 1
and 2 mm deep, with smoothly rounded edges. The trough
has been slightly distorted by lateral pressure, and would
have originally had the cross-section of a narrow steep-sided
‘V’. The trough walls were convex in section and the floor
was narrowly rounded. The trough is a narrow structure set in
a relatively broad buccal floor. The trough widens over the
posterior 12 mm of the symphysis, its floor becomes gently
concave, and its walls merge into the ventro-medial walls of
the jaw rami over the first 5 mm behind the symphysis.

A series of poorly preserved alveoli are visible on the
dorsal edge of each ramus, the more complete series on the
right extending back to about 50 mm behind the level of the
posterior end of the symphysis. They are small and ovoid,
between 1:5 and 2 mm labio-lingually and 3 mm antero-
posteriorly. The alveolar rims are not raised into cylindrical
collars like those of Plataleorhynchus, described below.
Alveolus 15 on the right side is exposed in section and is
deep, extending almost to the midline. Within the symphysis
is a median septum of bone separating the two series of
alveoli. Unlike the anterolaterally orientated spatula alveoli,
those of the post-spatula region are arranged perpendicular
to the midline, and the main tooth-row must have extending
out sideways from the jaw. The alveoli on the spatula and
anterior mandible were nearly horizontal, whereas those
further back were more dorsolaterally orientated, there being
a gradual gradation along the mandible. There are 29 alveoli
visible on the right dentary and about 30 were probably
present on each ramus (including the spatula teeth). They
imply the presence of a large number of small, outwardly-
directed teeth spaced evenly at approximately 3:25 mm
intervals.

SYSTEMATIC POSITION. SMC J5339 is clearly the mandible of
a ctenochasmatid pterodactyloid. It is shallow, has a long
symphysial region and would have borne about 30 laterally or
antero-laterally directed teeth of uniform cross-section on
each side (Fig. 2B). The small spatula would have borne ten
anterolaterally directed teeth. The buccal floor of the sym-
physial region has a broad flat surface curving down medially
to a narrow symphysial trough. Such flattened mandibles with
large numbers of evenly spaced, uniform, laterally orientated
teeth are only found in the Ctenochasmatidae, although the
symphysial trough appears at first to conflict with this posi-
tion as it has been used as a character to define the Orni-
thocheiridae.

The symphysial trough has previously been reported as an
ornithocheirid character (Wellnhofer 1978) but appears to
have a wider distribution, although in different configura-

_ tions. Ornithocheirids have either a shallow ‘U’-shaped sym-
| physial trough (Ornithocheirus sedgwicki and O. daviesi) or a

wide ‘V’-shaped trough (O. machaerorhynchus and Brasileo-

dactylus araripensis). In all these forms, the buccal floor
_ forms a narrow shelf on either side of the trough and the teeth
are vertically orientated. The holotype and only mandible
(BSPM AS VII 369) of the ctenochasmatid Gnathosaurus
subulatus is preserved with ventral aspect only exposed and
the presence of a trough cannot be established. However, the
palate of G. subulatus bears a low medial ridge presumably

developed from the co-ossified palatines. For the rostrum and

mandibles of G. subulatus to occlude in a useful manner,
| there must have been a corresponding medial trough in the
symphysial region of the mandibles of this species. The

palatal ridge, though low, is sharply keeled and narrow, and
the trough in SMC J5339 is of corresponding shape for such a
keel to fit into. This type of keel-trough system, in which a
small sharp keel and trough occur between wide palatal and
buccal shelves in flattened jaws, appears to characterise some
members of the Ctenochasmatidae. Thus the presence of a
symphysial trough does not characterise the Ornithocheiridae
sensu latu, but specific constructions of trough characterise
separately the Ornithocheiridae and some Ctenochasma-
tidae.

Within the Ctenochasmatidae, SMC J5339 is closest to
Gnathosaurus subulatus in configuration. The presence of 30
teeth per ramus is comparable to counts of 31 in G. subulatus
and ?36-38 in Plataleorhynchus, but distinct from 50-90 in
Ctenochasma and 25 in Huanhepterus. The small mandibular
spatula bearing 10 teeth is unlikely to have been the counter-
part to the large rounded rostral spatula bearing 22 teeth
found in Plataleorhynchus and the latter bears no medial
ridge that might correspond to the upper part of the ridge-
trough system in Gnathosaurus. In shape, the spatula of SMC
J5339 is comparable to that of G. subulatus and the latter
bears a medial ridge on the palate corresponding to the
symphysial trough of the mandible of SMC J5339. It is
proposed that SMC J5339 be placed in the genus Gnathosau-
rus as the species macrurus, distinguished from G. subulatus
by the difference in number of spatula teeth (10 in macrurus,
14 in subulatus) and the orientation of the post-spatula teeth
(lateral in macrurus, anterolateral in subulatus).

Genus PLATALEORHYNCHUS nov.

NAME. From the Greek: Platalea, the generic name of the
spoonbill, derived from Platys — flat; and Rynchos (latinised
to Rhynchus) — a snout.

TYPE SPECIES. Plataleorhynchus streptophorodon sp. nov.
DIAGNOsIs. As for the type and only species.

RANGE. Tithonian/Berriasian of the British Isles.

Figs 3-6

NAME. From the Greek: streptophoros, collar-wearing, and
odontos, a tooth.

Plataleorhynchus streptophorodon sp. nov.

DIAGNOSIS. Large ctenochasmatid attaining a skull length of
at least 400 mm, if Gnathosaurus-like proportions are
assumed. Spatula roughly circular and relatively sharply
demarcated from narrow rostrum posterior to it. Spatula
bearing 22 teeth with shallow roots not reaching to the centre
of the spatula. Anterior tip of palatines forming a slender
point wedged between the two premaxillaries. Palatal face of
spatula and anterior rostrum partly covered in rugose bone,
indicative of the presence of a keratinous pad. No ridge on
the midline of the palate. Over 62 (possibly up to 76) teeth
present on entire rostrum.

HOLOTYPE. BMNH R.11957. Rostrum of a pterodactyloid
visible in palatal aspect (Figs 3-5). Ex Corfe Castle Museum
collection, purchased by BM(NH) from Dorset County
Museum in 1958. Not previously described.

TYPE LOCALITY. Quarry near Langton Matravers, Purbeck,
Dorset. The precise locality is not known.

Horizon. ‘Middle Purbeck Beds’, Purbeck Limestone For-

mation; Tithonian, Upper Jurassic or Berriasian, Lower
Cretaceous. The precise horizon is not known.

DESCRIPTION. BMNH R.11957 is an incomplete section of a
pterodactyloid rostrum, 174 mm long, exposed in palatal
aspect, and comprising the premaxillaries, maxillaries and
palatines (Figs 3, 4A). These elements are partly fused and
only some sutures can be identified.

Spatula. The rostrum bears a terminal spatula which is
completely flat and shows no longitudinal or transverse
curvature (Fig. 5). It is formed by the premaxillaries which
are completely fused in the midline. It is 39 mm long and 32
mm wide and bears 22 alveoli around its periphery. The
anterior alveoli are damaged and represented by the broken
edges of the bases. The remaining alveoli on the spatula are
better preserved although the rims are never complete. The
spatula alveoli are between 0-75 and 1-25 mm apart and all
but the anterior pair are between 3-5 and 3-75 mm wide
labiolingually, the anterior pair being about half this size. The
alveoli in the anterior half of the spatula are anteriorly or
anterolaterally orientated, those in the posterior half gradu-
ally becoming more laterally orientated. The posteriormost
spatula teeth would have emerged posterolaterally from the
spatula, in sharp contrast to the anterior post-spatula teeth
which would have been laterally directed. In all cases, the
teeth would have emerged horizontally from the edges of the
jaws. The alveolar rims are formed from short raised collars
of bone, which tend to coalesce with neighbouring rims on
the spatula, giving it a slightly crenellated margin. Radiogra-
phy of the spatula has shown that the tooth sockets within it
are short and extend only partway to the middle of the
spatula. Although the teeth are missing, the small size of the
sockets suggests that the teeth themselves were not large or
required to cope with powerful movements of prey animals.
The palatal surface of the spatula is perforated by an outer
and an inner series of foramina (Fig. 5). The outer series
occurs in two parallel rows of elliptical foramina close to the
alveolar bases along the rostrum and forms a loop of 23 small
circular openings on the spatula. This series bears a one-to-
one relationship to the 22 alveoli (except for the third
alveolus on the left side which has 2) and the openings are
presumed to be the dental foramina. Each dental foramen is
set 4-5 mm in from the nearest tooth. The inner series of
foramina are larger, fewer in number (ten on the spatula) and

S.C.B. HOWSE and A.R. MILNER

more variable in shape. They are asymmetrically arranged
with three on the left side and seven on the right side. Several
emerge into shallow channels on the surface of the palate.
One pair of inner foramina at the ‘neck’ of the spatula
develop into diverging anterolaterally directed channels,
while the anterior pair of foramina on the spatula each
develop into forked anterolaterally directed channels sepa-
rated by a V-shaped notch.

The centre of the palatal surface of the spatula is covered
by a pad of rugose bone which puts out local anterolateral
extensions to the periphery of the spatula (Fig. 5). Posteriorly
the rugose patch becomes elongate and, at the level of the
‘neck’ of the spatula, it is flanked by two other elongate but
irregular rugose zones. Patches of rugose bone are also
present around and near the bases of the alveoli. The surface
of the central rugosity consists of a dense pattern of ovoid
cancellations separated by minute spans of bone. Near the
periphery of the spatula these cancellations become circular
pits. The non-rugose parts of the spatula are only lightly
pitted, as are the outer alveolar walls and most of the palatal
surface of the elongate region of the rostrum.

Post-spatula rostrum. Behind the spatula, the rostrum nar-
rows to 15 mm in width and then gradually broadens (Figs 3,
4A). The premaxillaries extend back behind the spatula for a
further 36 mm but here the medial suture is still present.
Behind the premaxillaries are the fused palatines in the
midline and the paired maxillaries on either side. The
palatines terminate in a narrow point anteriorly and are
slightly sloping to give a raised midline to the palate. The
maxillary-palatine sutures are largely visible, except where
damaged, but the premaxillary-maxillary sutures cannot be
seen clearly. The posterior palatal shelf of each maxillary
bears a long shallow trough between the alveoli and the
suture with the palatine. These troughs fade out well behind
the spatula. The damage to the rostrum appears to involve
some anteroposterior compression or telescoping with some
anteroposterior overlap of elements. The reconstruction of
the rostrum (Fig. 4B) attempts to compensate for this com-
pression, assumed to be equivalent to the space occupied by
3-4 teeth on each side.

There were at least 20 post-spatula teeth on each side and,

if it is assumed that some telescoping of the rostrum has |

occurred and that a few more teeth were present posteriorly,

Fig. 3 Plataleorhynchus streptophorodon gen. et sp. nov. BMNH R.11957, holotype rostrum in palatal aspect x 0-7.

E——EEE—————— EE

_THE PTERODACTYLOIDS FROM THE PURBECK LIMESTONE FORMATION OF DORSET

g-4 Plataleorhynchus streptophorodon gen. et sp. nov. BMNH R.11957, holotype rostrum in palatal aspect. (A) Interpretive drawing. (B)
Reconstruction of rostrum. The reconstruction incorporates compensation for the telescoping of the original specimen, and is therefore
somewhat longer. Abbreviations: al. = alveolus; d.f. = dental foramen; MAX = Maxillary; PAL = Palatine; PMX = Premaxillary; v.f. =
' vascular foramen; x—x crack marking region of telescoping of specimen. Scale = 10 mm.

|
|

there might have been 5-7 more in each row (Fig. 4B
reconstruction). Their alveoli are similar in size to those of
the spatula teeth, but are less closely spaced with 2-5 to 3-5
mm separation, and their ‘collars’ show no tendency to
coalesce. The dental foramina move to close proximity to the
teeth behind the spatula and are incorporated into the base of
the alveolar rim by tooth 17 on each side. On either side of
the midline of the post-spatula rostrum are a series of large
oval foramina, corresponding to the inner foramina on the
spatula. Most of the palatal surface in this region is covered in
delicate anteroposteriorly orientated striae, with occasional
rugose patches along the midline. The rugose palate surface.
In several groups of vertebrates, notably chelonians, birds
and some ornithischian dinosaurs, the presence of a horny
beak or palate is associated with a characteristic rugose
surface to the bone underlying the horn-covered area. This is
noticeable in the palates of many living turtles and birds,
where the association between the horny beak and crushing
palate and the rugose and vascular underlying bone is indis-
putable. Such palatal surfaces have not been previously
reported in tooth-bearing pterosaurs, but are present in
Plataleorhynchus as described here. A search through speci-
mens available to us has revealed a slender strip of rugose
bone on the anterior palate of the anhanguerid ‘Ornithochei-
rus’ cuvieri from the Lower Chalk of Kent (pers. obs.) and it
is likely that such structures were present in other large
pterodactyloids at least. It appears that Plataleorhynchus had
a pad of horny skin on the palatal surface of the spatula and
along parts of the anterior palatal surface of the rostrum.
Such horny palatal surfaces generally occur where a hard
renewable surface is required by a feeding technique involv-
ing crushing or abrasion, which would cause damage to a
surface that was not both hard and self-regenerating.

SYSTEMATIC POSITION. The only pterosaurs to have a ros-
trum with such a pronounced flat spatula and slender, later-
ally directed teeth are some members of the family
Ctenochasmatidae, also known from the Upper Jurassic of
Europe and China. Plataleorhynchus can be distinguished
from each of the three previously described ctenochasmatids
as follows:

Ctenochasma Meyer, 1851, represented by three species from
Germany and France, is the type genus of the family. It is
known only from relatively small specimens, lacks a spatula
and has a much larger number of closely spaced teeth
(100-180) in the upper jaw. Although the only rostrum of
Plataleorhynchus is incomplete and the full rostral tooth
count is unknown, it is unlikely to be much greater than 76.

Huanhepterus Dong, 1982, represented by the single species
H. quingyangensis from China, is of suitable size, has a
terminal expansion to the rostrum but is estimated to have a
rostral tooth count of about 50, significantly fewer than the
other ctenochasmatid genera including Plataleorhynchus.

Gnathosaurus Meyer, 1834, represented by G. subulatus
from Germany and G. macrurus from Purbeck has a similar
number of rostral teeth to Plataleorhynchus, but the spatula is
less circular, more ovoid in ventral aspect, bears only 10-14
teeth, and the tooth orientation change between the posterior
spatula dentition and the anterior post-spatula dentition is
gradual. The anterior region of the spatula bears very large
teeth, the roots of which extend into the middle of the
spatula. There is no evidence of rugosity on the palatal

S.C.B. HOWSE and A.R. MILNER

surface of the spatula or rostrum. In Plataleorhynchus, the
spatula is almost circular in plan view, bears 22 teeth and the
alveoli show a very sharp demarcation in orientation at the
spatula-postspatula boundary. The anterior marginal teeth of
the spatula do not appear to have been large and their sockets
extend only partway towards the middle of the spatula. The
palatal face of the spatula and anterior rostrum are partly
rugose, as described above.

Plataleorhynchus is closest to Gnathosaurus in general
configuration but is larger and has a rostrum showing greater
differentiation between the spatula and post-spatula areas. In
conclusion, Plataleorhynchus can be argued to represent a |
new genus of large ctenochasmatid with a large spoonbill-like
rostrum.

LIFE STYLE. Most ctenochasmatids appear to have been
filter-feeders, the long rostrum bearing long laterally’
directed, intermeshing teeth, providing a sieve-like device for
filtering small crustaceans or similar organisms from the
water or mud. Plataleorhynchus may likewise have been a
sweep-feeder like the living spoonbill Platalea, swinging the
head from side to side and filter feeding on a large scale. It is
possible that Plataleorhynchus may have fed in a slightly
different manner, in that, compared to other large ctenochas- |
matids, the terminal spatula was more differentiated, the)
teeth were relatively smaller (to judge from the alveoli) and
the palatal surface appears to have been covered by a horny|
surface (Fig. 6). These suggest that the animal may have used _
its spatula and anterior dentition to rake through mud or)
weed to disturb animals, then gripping and crushing slightly
larger animals in the water or substratum with the horny
spatula as they were disturbed. This may have been a more)
significant means of procuring food for this large ctenochas-
matid than simply filtering water with organisms in it.

CTENOCHASMATIDAE Incertae sedis

CERVICAL VERTEBRAE. SMC J5340 is a middle series cervi-
cal vertebra of a pterodactyloid, exposed in dorsal aspect and
dorso-ventrally crushed. The specimen was figured by Seeley
(1875 fig.1 (unnumbered), 1901 fig.65) and Howse (1986 figs) «:
8-9). It came from near Langton Matravers, Purbeck, Dorset ©
(Seeley 1875) and probably from the same horizon as thi
lectotype of Gnathosaurus macrurus. The precise locality 1
not known.

SMC J5340 is either a fourth or, more probably, a fift
cervical vertebra of a long-necked pterodactyloid. It was mos
recently redescribed by Howse (1986) and the following is
brief summary of that description. The specimen has bee
crushed in a generally dorso-ventral plane with some latera
distortion, so that the apex of the neural arch has bee
displaced to the left of the centrum midline. The centrum is .,
109 mm long, but is largely obscured by the neural arch. The > )
anterior cotyle and posterior condyle of the centrum are too,
crushed for much of their structure to be made out, although .
the posterior condyle was situated between a pair of postex
apophyses, the left one of which is preserved. There is a smal
oval pneumatic foramen about halfway along the centrum
visible on the exposed right side.

The neural arch appears to have been a low-vaulted
somewhat rounded structure. It is broad-ended, tapering to¢ ,_
narrow waist about two-thirds of its length from the anterior)
The anterior margins of the neural arch were extended int

THE PTERODACTYLOIDS FROM THE PURBECK LIMESTONE FORMATION OF DORSET 81

4 an
6
° “48

ne ‘y

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ire
°
Ay
UHC}
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aetin)
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Abbreviations: al. = alveolus; d.f. = dental foramen; v.f. = vascular foramen. Scale = 10 mm.

in 5 Plataleorhynchus streptophorodon gen. et sp. nov. BMNH R.11957, spatula of rostrum in palatal aspect (interpretive drawing).
|
|

darallel, elongate, horn-like prezygapophyses, the left one of

which is preserved. It bears an ovoid articular facet. The

dostzygapophyses, represented by a worn left structure, were
yroad blunt processes each bearing a distinct tubercle. The

ieural spine is a low slender ridge of bone running the full
ength of the vertebra. Fracturing at both ends suggests that it
nay have been slightly more raised in these regions.
BMNH 48387 (Fig. 7) is a second, incomplete, cervical
vertebra which is part of the Beckles Collection, purchased in
$877. It is recorded merely as having been collected from the
/Middle Purbeck Beds’ of Durlston Bay. It is a fragment
‘omprising the posterior end of a flattened and elongate
ervical vertebra similar to the above specimen. Like SMC
5340, it has a low neural spine and bears a pair of postexapo-
hyses which bracket the posterior condyle.

These elongate cervical vertebrae clearly do not belong to
n ornithocheirid, a gallodactylid or some pterodactylids.
uch vertebrae are found in the Ctenochasmatidae, the
zhdarchidae and some species of Pterodactylus, including
*. antiquus and P. longicollum (the latter two possibly

synonymous according to Bennett (1993)). In recent work
SMC J5340 has been treated as a fragment of a possible early
azhdarchid pterodactyloid (Howse 1986, Wellnhofer 1991a).
However, now that it has been established that two suitably
large ctenochasmatids, Plataleorhynchus and Gnathosaurus,
are present in the Purbeck Limestone Formation, the system-
atic position of this cervical must be reconsidered. The larger
ctenochasmatid Huanhepterus has cervicals of similar size and
proportion and this size and shape of cervical vertebra could
reasonably be predicted to be present in Plataleorhynchus.
Nevertheless, given the diversity of long-necked forms in the
slightly earlier pterodactyloid fauna from the Solnhofen
Limestone, such a cervical cannot be confidently assigned to
a particular genus, and it is here treated as indeterminate
ctenochasmatid material.

82 S.C.B. HOWSE and A.R. MILNER

l S.C.8. HOWSE

Fig. 6 Restoration of the head of Plataleorhynchus streptophorodon, to show relationship of spatula to the head, using the ctenochasmatids
Gnathosaurus and Ctenochasma as models.

Fig. 7 Incomplete indeterminate ctenochasmatid cervical vertebra. BMNH 48387. Abbreviations: n.s. = neural spine; px. =
postexapophysis; pz. = postzygapophysis. Scale = 10 mm.

THE PTERODACTYLOIDS FROM THE PURBECK LIMESTONE FORMATION OF DORSET

Family ORNITHOCHEIRIDAE Seeley, 1870

SYSTEMATIC NOTE. Bennett (1989) has proposed uniting a
range of advanced pterodactyloids including the orni-
thocheirids, pteranodontids and ‘Ornithodesmus’ in one fam-
ily under the name Pteranodontidae Marsh, 1876. In this
work, we follow the practice of Wellnhofer (1991b) in divid-
ing these forms into several families, pending a more compre-
hensive analysis of their relationships.

Genus ORNITHOCHEIRUS Seeley, 1869

TYPE SPECIES. Ornithocheirus compressirostris (Owen) See-
ley: from the Turonian of Burham, Kent, England (see
_Wellnhofer 1978 pp. 55-56 for discussion of the identity of
_ the type species of Ornithocheirus).

‘Ornithocheirus’ sp. A Fig. 8

/ MATERIAL. BMNH R.11958, the anterior region of a man-
| dible of a pterodactyloid exposed in ventral aspect (Fig. 8).
|The specimen is somewhat crushed and the teeth have all
' been lost. Ex Corfe Castle Museum collection, purchased by
' BM(NH) from Dorset County Museum in 1958.

TYPE LOCALITY. ‘Near Swanage’, Dorset. The precise local-
' ity is not known.

HORIZON. ‘Middle Purbeck Beds’, Purbeck Limestone For-
mation; Tithonian, Upper Jurassic or Berriasian, Lower
Cretaceous. The precise horizon is not known.

DESCRIPTION. BMNH_ R.11958 comprises the anterior
‘tegion of a pterodactyloid mandible exposed in ventral aspect
on a block of limestone (Fig. 8). It consists of a complete
‘elongate symphysial region and posteriorly incomplete rami,
the length of the incomplete specimen being 208 mm in the
midline. The symphysial region is 118 mm long and relatively
Narrow, passing from 9 mm width near the tip, to 20 mm
width where the rami diverge. There is a slight spatulate
expansion of the tip. A short section of the symphysis is
visible in dorsal aspect and shows the presence of a median
trough with a deep ‘V’-shaped cross-section, and a few oval
alveoli. The buccal shelves on either side of the trough are
relatively narrow.

Most of the alveoli of the symphysial region cannot be
»bserved directly although their lateral edges can be seen as
indulations along the jaw margin, at least demonstrating that
eeth were present in this region. The few alveoli visible
\ppear to be widely spaced at intervals of about 6 mm. If this
s uniform, there would be space for about 23 on each ramus.
‘he ventral midline of the symphysial region bears a distinc-
ive low keel. For the first 26 mm of its length, the symphysis
_5 a flattened structure and the keel, which originates 9 mm

ehind the tip, is a low slender ridge about a millimetre wide

d with a semicircular cross-section. For the next 35 mm, the

mphysial region becomes a deeper, steeper-sided structure

nd the keel is a deeper ridge with a V-section merging into
le sides of the mandible. For the posterior 56 mm of its
ngth, the symphysial region is severely crushed, but the
lore resistant keel has resisted the crushing and forms a
arp crest extending to the back of the symphysis.
The rami are crushed outwards so that their width of 5-6
m represents their original depth. A consequence of this
ushing is that the dorsal edges face laterally and it can be

Fig. 8 ‘Ornithocheirus’ species A. BMNH R.11958, anterior region
of mandible in ventral aspect x 0-8.

seen that there are no alveoli behind the level of the
symphysis. Crushing has splayed the rami slightly and they
probably diverged less in life. The rami are broken posteri-
orly, anterior to the position of the articulars.

SYSTEMATIC POSITION. The discovery and description of a
series of ornithocheirids, criorhynchids and anhanguerids
from the Santana Formation of Brazil has resulted in the
systematics of this group passing though a period of instability
while new relationships and systematic criteria are defined.
The Ornithocheiridae as used in recent years appears to be
defined on negative criteria, and represents those members of
the old Ornithocheiridae which lack the cranial outgrowths of
the criorhynchids and anhanguerids. The pterodactyloid
mandible described above exhibits two systematically signifi-
cant characters, namely the presence of a ‘V’-shaped trough
extending along the midline of a symphysis, and relatively
widely spaced oval alveoli. The only genera possessing these
features, and an associated palatal ridge, are those until
recently placed in the Ornithocheiridae sensu latu, but now
distributed among several closely related families including
the Criorhynchidae and the Anhangueridae (see Wellnhofer
1991b for a review of the taxonomy of the Brazilian members
of this group). The palatal ridge and symphysial trough are
known only in the Ornithocheiridae and Ctenochasmatidae
and have not been described in any other Jurassic pterodacty-
loids, nor in Dsungaripterus, Ornithodesmus, Noripterus,
Pteranodon or Nyctosaurus.

In the following systematic discussion, comparison is made
with mandibular material of the following (sources quoted in
parenthesis):

Ornithocheiridae Seeley, 1870:

‘Ornithocheirus’ [Cambridge Greensand] (personal obser-
vation)

Brasileodactylus [Santana Formation] (Kellner 1984).
Anhangueridae Campos and Kellner 1985:

Anhanguera [Santana Formation] (Wellnhofer 1991b)
Criorhynchidae Wellnhofer 1991b:

Tropeognathus [Santana Formation] (Wellnhofer 1987).

Ornithocheirus species A, represented by BMNH R.11958,
has a very slight spatulate expansion, a long parallel-sided
symphysis bearing a deep ‘V’-shaped symphysial trough, a
very low ventral V-section keel and a complete absence of
ventral blades or crests. The Anhangueridae are character-
ized by blades and crests on the rostrum and mandible and
the mandible bears a shallow ‘U’-shaped symphysial trough.
BMNH R.11958 is therefore not an anhanguerid. The Crio-
rhynchidae are characterized by a massive blade of bone on
the anteroventral end of the mandible and it is therefore not a
criorhynchid, although the ‘V’-shaped symphysial trough
does occur in that family. The Ornithocheiridae is at present
a residue comprising the crestless members of the old Orni-
thocheiridae sensu latu. Within the Ornithocheiridae, BMNH
R.11958 most closely resembles some of the mandibular
material from the Cambridge Greensand. The taxonomic
status of this material has not yet been rationalized with
respect to the Santana Formation pterodactyloids (that work
is being undertaken by Dr D. Unwin, University of Bristol),
and therefore the nomenclature of Seeley and Hooley is used
in the following discussion. The Greensand specimen that
most closely resembles BMNH R.11958 is the type of Orni-
thocheirus machaerorhynchus Seeley, 1870 (pl.xii, figs 1, 2), a
fragment from the distal region of a mandibular symphysis

S.C.B. HOWSE and A.R. MILNER

dist.

Fig. 9 Doratorhynchus validus (Owen) Seeley, nomen vanum.

BMNH 40653, holotype phalanx of digit IV of right manus.
Abbreviations: dist. = distal; preax. = preaxial; prox. =
proximal. Scale = 10 mm.

THE PTERODACTYLOIDS FROM THE PURBECK LIMESTONE FORMATION OF DORSET

broken at both ends. The specimen is derived from a deep
symphysial region, acutely triangular in cross-section and
with a ventral keel. The alveoli are oval in cross section and
evenly spaced. The buccal shelves on either side of the
symphysial trough are narrow, and the trough itself is deep
and ‘V’-shaped in cross section. No other mandibular mate-
rial from the Cambridge Greensand has precisely this cross-
section, but some rostral fragments bear a ‘V’-shaped medial
ridge on the palatal surface which appears to complement the
trough in O. machaerorhynchus. Figured examples of such
rostral fragments are the type specimens of O. denticulatus
Seeley, 1870 (pl.xui, figs 8-9) and O. microdon Seeley, 1870
(pl.xu figs 6-7). Other comparable specimens described by
Seeley (1870) but not figured include the type rostral frag-
ments of O. polyodon, O. nasutus, O. dentatus, O. oweni and
O. tenuirostris. Three specimens, the types of O. machaero-
rhynchus, O. oweni and O. tenuirostris were transferred by
Hooley (1914) to the genus Lonchodectes while the rest were
_ retained in Ornithocheirus.
Other Greensand and Chalk ‘ornithocheirids’ including O.
- daviesi, O. sedgwicki, O. cuvieri, O. scaphorhynchus, O.
» platysomus and O. brachyrhinus, have a shallow palatal ridge
and a shallow ‘U’-shaped glossal trough, quite distinct from
_ those described above. The shallow type of trough also occurs
in the Anhangueridae (but not the Criorhynchidae) and it is
; possible that this suite of specimens are anhanguerids.
BMNH R.11958 is significant because of its early age, but
) isnot amenable to precise systematic placement. The absence
| of crests and other outgrowths, the widely spaced vertical
_ teeth and the shape of the symphysial trough all permit it to
_ be associated with certain of the Greensand Ornithocheirus
‘species’ of Seeley, all of which were placed by Hooley (1914)
‘in either Ornithocheirus or Lonchodectes. It thus appears to
_be a member of the Ornithocheiridae sensu stricto and not an
janhanguerid or a criorhynchid. It must be attributed to
| Ornithocheirus pending a satisfactory resolution of the sys-
| tematics of that genus. Most of the named species already in
‘the literature which incorporate or are based on comparable
elements are nomina vana restricted to type specimens which
|are mandibular fragments (e.g. ‘O.’ machaerorhynchus). In
‘the absence of any unique diagnostic characteristics, BMNH
'R.11958 cannot be the basis of a new species despite its
distinct stratigraphical position, and it would be misleading to
refer it to O. machaerorhynchus on the basis of the limited
similarity of mandibular fragments. Pending revision of the
Greensand material, the simplest and most justifiable proce-
fae is to record the specimen as Ornithocheirus ‘species A’.

| PTERODACTYLOIDEA Incertae sedis

_Doratorhynchus validus (Owen) Seeley nomen
vanum Fig. 9

870 Pterodactylus validus Owen: pl.XIX, fig.7.

875 Doratorhynchus validus (Owen) Seeley: 465-8 partim.

888 © Ornithochirus validus (Owen) Lydekker: 26-7.

890 Doratorhynchus validus (Owen); Woodward & Sher-
born: 227 partim.

929 Doratorhynchus validus (Owen); Plieninger: 27-28

| partim.

958 Ornithocheirus validus (Owen); Delair: 71-2.

és Doratorhynchus validus (Owen); Wellnhofer: 58
partim.

ot set
A
tae a
1g '
+
f
1: ,
1
ae:
ie i;
(ee ‘
la 7
ht .
iY y
ae)
if H
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‘3
a
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i
By
ey
ai

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A prox.

Fig. 10 Pterodactyloidea incertae sedis. A) BMNH 2462 right
radius in ventral aspect. B) BMNH R.5798 left tibia in posterior
aspect. Abbreviations as for figure 9. Scale bars = 10 mm.

The above synonymy is restricted to publications which make
specific reference to the holotype phalanx.

HoLotyPeE. BMNH 40653, a first or second phalanx of digit
IV (the wing-finger) of the right manus of a large pterodacty-
loid (Fig. 9). The specimen is exposed in dorsal aspect and
was figured previously by Owen (1870 PI.XIX, fig.7).

LOCALITY. Simply noted by Owen 1870 as ‘Swanage’, but
probably from the Langton Matravers quarry.

Horizon. ‘Middle Purbeck Beds’, Purbeck Limestone For-
mation; Tithonian, Upper Jurassic or Berriasian, Lower
Cretaceous.

DESCRIPTION. BMNH 40653 is a crushed phalanx of digit IV
(the wing-finger) of a pterodactyloid (Fig. 9). It is about 300
mm in length, with the proximal epiphysis 31 mm across and
the distal epiphysis 19 mm across. There is one transverse
fracture near the middle of the shaft and some superficial
longitudinal fractures. The leading edge of the phalanx is
straight, while the trailing edge is largely straight but curves
back at each end into the posterior margins of the epiphyses.
The exposed face of the bone appears to have been flat, prior
to crushing, rather than of convex section, and thus probably
represents the dorsal face of the phalanx. If so this was a
phalanx of the right manus. The specimen was identified by
Owen as the second phalanx of digit IV, and if so, would have
been part of a wing finger of 1-2-1-5 metres in length. The
size and extent of curvature of the distal end indicate that it
was either a first or second phalanx. The shape of the
proximal end resembles that of a first phalanx with the
extensor process broken off. The specimen could be either a
first or second phalanx of digit IV and is clearly not determi-
nate below subordinal level. Thus the binomen Doratorhyn-
chus validus is anomen vanum restricted to the type specimen
and no other Purbeck material can be referred to it, particu-
larly as there are clearly at least three pterodactyloids in this
assemblage.

OTHER PTERODACTYLOID MATERIAL
FROM PURBECK

The following anatomically determinate pterodactyloid speci-
mens have also been collected from the Purbeck Limestone
Formation. One was recorded in Lydekker’s (1888) cata-
logue, but several have not been mentioned in the literature.

TEETH. Ensom, Evans & Milner (1991) have reported a new
microvertebrate assemblage from Sunnydown Quarry, Sut-
tle’s Quarry and other localities near Langton Matravers,
Purbeck. This material includes several slender recurved
pterosaur teeth (DORCM), which Ensom et al. identified as
thamphorhynchoid teeth, assuming them to represent the
anterior fish-spearing teeth of small rhamphorhynchoids. An
alternative interpretation is that they are the slender recurved
marginal teeth of a large ctenochasmatid such as Gnathosau-
rus or Plataleorhynchus. This cannot yet be demonstrated as
no intact teeth have been found with either specimen, but this
would represent the most conservative interpretation of the
systematic position of these teeth.

RADIUS. BMNH 2462 (Fig. 10A) is a small pterodactyloid
radius from ‘the Middle Purbeck of Swanage’, originally in

S.C.B. HOWSE and A.R. MILNER

the G. A. Mantell Collection and acquired by the BM(NH) in
1838. It was listed as a phalanx by Lydekker (1888 p.27),
mentioned by Wellnhofer (1978) and first identified as a
radius in 1988 by S.C. Bennett (Kansas) during study of the
BM(NH) collections. The radius appears to be a right ele- |
ment exposed in ventral aspect. It is 111 mm long, 13 mm
across the proximal epiphysis, 11 mm across the distal epiphy- |.
sis and 4 mm across the diaphysis at its narrowest diameter. |
Beyond being recognisable as a pterodactyloid radius, it is)
indeterminate.

METACARPAL. BMNH 48380 (Fig. 11) is a fourth metacarpal
of a medium-size pterodactyloid. It formed part of the S. H.
Beckles Collection and is recorded as being from the ‘Middle
Purbeck Beds’ of Durlston Bay. It is incomplete, lacking the’
proximal end, but the distal end and the main shaft are about ‘F
44 mm in length, and the entire bone would have been 50-55
mm long. It bears the characteristic distal paired roller-joint’_
articulation with the first phalanx of digit 1V. The exposed)
semicircular articulation is smaller than the largely buried one
below it, traces of which are visible. This suggests that the
specimen is exposed in ventral aspect, the ventral articulation
being the smaller one.

PHALANGES. DORCM G.100 is a phalanx from the ‘Middle) ”
Purbeck Beds’ of Langton Matravers originally belonging to
the J. C. Mansell-Pleydell Collection. It was listed by Delair
(1958 p.71), but is determinable only as a pterodactyloid| ~
phalanx.

BMNH 48387 is the register number for a collection of |
phalangeal fragments from the S. H. Beckles Collection, all"
originating from the ‘Middle Purbeck’ of Durlston Bay. Most
are indeterminate, but one represents the proximal region of
phalanx 1 of the wing finger — digit IV. It bears the character
istic cup-shaped proximal articulation formed partly by the
extensor process.

TiBIA. BMNH R.5798 (Fig. 10B) is a large and very slender

tibia collected from the Swanage area by S.L. Wood in 1919
and originally identified as a phalanx. It is 205 mm long an¢
poorly preserved but appears to be a left tibia visible 1
posterior aspect. It must have belonged to a large, long) ..
legged pterodactyloid, and may well be a ctenochasmatic
tibia.

[
DISCUSSION i
| Ee
Chronological ranges of pterodactyloid families ‘a
The recognition of ctenochasmatids and ornithocheirids it
the Tithonian-Berriasian of England does not significantly
change our perception of the geographical distribution 4
these forms but does extend their known chronologica
ranges, and also alters the probable range of the Azh
darchidae.

The Ctenochasmatidae were already known from th
Upper Jurassic of Europe. The Purbeck Plataleorhynchu
and Gnathosaurus do not represent an extension to thei
known geographical distribution but may represent a sligh |
chronological extension up into the earliest Cretaceous.

The Azhdarchidae, previously suggested to extend downt
the base of the Cretaceous, on the basis of the cervice
vertebra from Purbeck, may not appear until later in t
record. The discovery of Plataleorhynchus does demonstrat

THE PTERODACTYLOIDS FROM THE PURBECK LIMESTONE FORMATION OF DORSET 87

that ctenochasmatids as large as the Chinese Huanhepterus
were present in Europe. Huanhepterus is remarkable for the
extreme elongation of the middle series cervical vertebrae
and it is probable, though not demonstrable, that the long
cervical vertebra from Purbeck belongs to either Gnathosau-
rus or Plataleorhynchus and that members of both of these
genera had long Huanhepterus-like necks. Howse (1986)
suggested that the long cervical SMC J5340 from Purbeck
represented an early record of an azhdarchid pterodactyloid,
and that ctenochasmatids were relatives of the azhdarchids,
sharing neck elongation but exhibiting it to a lesser degree.
An alternative possibility is that SMC J5340 and BMNH
48387 belonged to a large ctenochasmatid and that the
ctenochasmatids had entirely similar cervical series to the
azhdarchids which may or may not have been present at the
Jurassic-Cretaceous boundary.

The Ornithocheiridae have been described from several
localities in the Cretaceous of England, the previous earliest
record being O. sagittirostris from the Valanginian Hastings
Sand of St Leonards on Sea, Sussex. The occurrence of an
ornithocheirid in Tithonian-Berriasian strata is a significantly
earlier record, and shows that the group was present in the
uppermost Jurassic. This implies that ornithocheirids were
present at the time that the Solnhofen Limestone was being
deposited, and the possibility that one or more of the many
Solnhofen pterodactyloids may be primitive or juvenile orni-
thocheirids should be explored.

Cladograms of possible pterodactyloid relationships published
by Howse (1986 fig.11) and Bennett (1989 fig.3) share the
conclusion that most of the higher pterodactyloids can be
grouped in two major clades — the ‘long-spined’ forms with tall
cervical neural spines (Ornithocheiridae and Pteranodontidae),
and ‘long-necked’ forms (some Pterodactylidae, Ctenochasma-

| tidae, Pterodaustridae and Azhdarchidae). Long-necked forms
| have been known from the Tithonian Solnhofen Limestone since
Pterodactylus antiquus was first described in 1812, but the new
Purbeck material confirms that the presumed ‘long-spined’
sister-group was also present at the Tithonian-Berriasian bound-
ary. If Bennett’s cladogram is accepted, this would also imply that
those pterodactyloid families which branched off further down
the cladogram — the Nyctosauridae and the Dsungaripteridae —
were also present in the late Jurassic, though they are known only
‘from Cretaceous specimens at present. The relationships of
)pterodactyloids are not yet so robustly established that we can
draw firm conclusions about their phylogeny. However, it
‘increasingly appears that the major family-level groups of
pterodactyloids had diversified by the beginning of the Creta-
ceous although most are not represented until at least the
Valanginian.

The Purbeck vertebrate fauna

The Purbeck Limestone Formation produced one of the first
diverse assemblages of mid-Mesozoic continental vertebrates
to be collected in the mid-19th century, principally from the
Beckles Pit and Durlston Bay, and largely comprised of
lizards, crocodiles, turtles, mammals and a few dinosaurs.
For much of this century, little new material was collected
and the known tetrapod fauna of the Purbeck Limestone
Formation has remained relatively unaltered, subject to
inevitable taxonomic revisions from time to time. However,
recent systematic collecting of microvertebrate material from
quarries around Langton Matravers by P.C. Ensom, and a
series of fortuitous discoveries in the field and in museums, is
adding numerous new forms to this Tithonian-Berriasian
continental fauna.

The new microvertebrate assemblages collected by Ensom
have proved to contain four species of lissamphibian compris-
ing a frog, two types of salamander (one a batrachosauroi-
did), and an albanerpetontid (Ensom, Evans & Milner 1991),
together with three new genera of multituberculate mammal
(Kielan-Jaworowska & Ensom 1992; 1994). A fortuitous
discovery in Durlston Bay has added the large sphenodontid
Opisthias to the Purbeck fauna (Evans & Fraser 1994), and
reassessment of older material has added the sphenodontid
Homoeosaurus (Whiteside 1986), a new anguimorph lizard
(Evans, 1994) and the pterodactyloids described in this work.

As a result of these and potential further additions, the
Purbeck fauna is becoming one of the richest mid-Mesozoic
continental tetrapod assemblages known, comparable to
those of the Late Cretaceous of North America and Mongo-
lia. Given that the assemblage has been collected from a
restricted range of strata over a few square miles, it is likely to
represent a genuine fauna of coexisting vertebrates and
consequently has considerable potential for palaeoecological
studies. It has always been clear that there was a substantial
freshwater component to the Purbeck fauna, as evidenced by
the presence of a diversity of turtles and crocodiles. The
discovery of a range of lissamphibians, together with two
pterodactyls specialised for feeding on small aquatic organ-
isms, further emphasises the freshwater-marginal source of
much of the fauna.

ACKNOWLEDGEMENTS. We thank Dr Angela C. Milner (Natural
History Museum, London), Dr David B. Norman (Sedgwick
Museum, Cambridge) and Dr Peter Wellnhofer (Bayerische Staat-
sammlung ftir Palaontologie und Historische Geologie, Munich) for
permission to study material in the collections in their care. Our
particular thanks go to William Lindsay (Natural History Museum)

jig. 11 Pterodactyloidea incertae sedis. BMNH 48380 fourth metacarpal in ventral aspect. Abbreviations as for figure 9. Scale = 10 mm.

for his meticulous preparation of the new specimens. Our work on
this material benefited greatly from discussions with Dr Wellnhofer
and the typescript also benefited from comments from Mr Paul C.
Ensom (York Museum). The photographs were provided by the
Photographic Unit of the Natural History Museum, London. This
work was funded by the Natural Environment Research Council

under Grant GR3/8053.

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— 1875. On an Ornithosaurian (Doratorhynchus validus) from the Purbeck
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— 1901. Dragons of the Air. An Account of Extinct Flying Reptiles. Methuen
and Co., London. 239 pp.

Wellnhofer, P. 1978. Pterosauria. In Wellnhofer, P. (ed.). Handbuch der
Paldoherpetologie, 19. 82 pp. Gustav Fischer Verlag, Stuttgart.

—— 1987. New crested pterosaurs from the Lower Cretaceous of Brazil.
Mitteilungen Bayerische Staatssammlung fiir Paldontologie und historische
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—— 1991a. The illustrated encyclopedia of pterosaurs. Salamander Books,
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—— 1991b. Weitere Pterosaurierfunde aus der Santana-Formation (Apt) der
Chapada do Araripe, Brasilien. Palaeontographica Abteilung A, 215:
43-101.

Whiteside, D.I. 1986. The head skeleton of the Rhaetian sphenodontid
Diphyodontosaurus avonis gen. et sp. nov. and the modernizing of a living
fossil. Philosophical Transactions of the Royal Society, London Series B, 312:
379-430.

Woods, H. 1891. Catalogue of the type fossils in the Woodwardian Museum,
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Woodward, A.S. & Sherborn, C.D. 1890. A Catalogue of British Fossil
Vertebrata. Dulay and Co., London. 396 pp.

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CONTENTS

1 Asynopsis of neuropteroid foliage from the Carboniferous and Lower Permian of Europe
C.J. Cleal and C.H. Shute
53 The Upper Cretaceous ammonite Pseudaspidoceras Hyatt, 1903, in north-eastern Nigeria
P.M.P. Zaborski
73 The pterodactyloids from the Purbeck Limestone Formation of Dorset
S.C.B. Howse and A.R. Milner

Bulietin of The Natural History Museum
GEOLOGY SERIES
Vol. 51, No. 1, June 1995

SAM |1OA0

Geology Series

MUSEUM

VOLUME 51 NUMBER 2 30 NOVEMBER 1995

The Geology Series is edited in the Museum’s Department of Palaeontology
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Geology Series
ISSN 0968-0462 Vol. 51, No. 2, pp. 89-306

The Natural History Museum
Cromwell Road
London SW7 5BD Issued 30 November 1995

Typeset by Ann Buchan (Typesetters), Middlesex
Printed in Great Britain at The Alden Press, Oxford

Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):89

Palaeontology of the Qahlah and Simsima

Formations (Cretaceous, Late

Campanian-Maastrichtian) of the United

Arab Emirates-Oman Border Region

CONTENTS

Preface MaKe Howarth (6diton)) i ciiee ce cnecacese esas ceceiepan sansa tacauen aray demsadsagitar ces seaemunaaeceesakis

Late Cretaceous carbonate platform faunas of the United Arab Emirates-Oman border region

A.B. Smith, N.J. Morris, W.J. Kennedy and A.S. Gale ........... 0. cece cece cece ee eee ee eeneeeeeeneneeeees

Late Campanian-Maastrichtian echinoids from the United Arab Emirates-Oman border region

PACAES STM pes cece = se occa Sa ciasis lessees sid eineier posse eee ety eGhecieaaasie seicasie sions able'aisslssowsiegietysieseaaseseslancs

Maastrichtian ammonites from the United Arab Emirates-Oman border region

WAU peING ING Uiyare tee ee acne nen aoc oat sini sais stersiast nye ratiiscuberarannsisioseile me sislbitn adelaatins oajlnioccleilelseteclncle.teaielpbtte wel

Maastrichtian nautiloids from the United Arab Emirates-Oman border region

IN| UM Woy tit Gh. arene tospbatoestuddonne boas bebe dodpono sun nacaue oounqHdecoUCdelico sucocrae bocsuccr enc encneeenncUMaea serene ness

Maastrichtian Inoceramidae from the United Arab Emirates-Oman border region

INiVJo IIOAS’ Sabnodecesubandoa sootoresndoondoddtSaeaaodsessbesuacosenaadacccodeceda1 Suuaadet an SaUnUedHPbEsonRBpeenree anes

Late Campanian-Maastrichtian Bryozoa from the United Arab Emirates-Oman border region

12,1 Ds SAH Ot re osadaaabaabanden bacAdqacsnonte Sacer ap baad nEeren be poon An icapncacotnoar anaMe tne aneuRMaceaAconaacacose: maces

Maastrichtian Brachiopods from the United Arab Emirates-Oman border region

IIR (OMEN -docsssondensous goude cba so eonaoEtbooEndeacoebeoaceotb eae say menbodadone Gad ASaeRUe RD oos noe aac BoC racnEcrcanes

Late Campanian-Maastrichtian rudists from the United Arab Emirates-Oman border region

NET MOrnissanduesWwriskeltons ca-ctccscacsossceceimaecnasars soa eoacoca te ascionue cence ease cehiilecmstoutsene es

(© The Natural History Museum, 1995
|

Issued 30 November 1995

Sak een eMac 275

Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):90

Preface

M.K. HOWARTH

Department of Palaeontology, The Natural History Museum, London SW7 5BD

The eight papers in this volume describe the Upper Creta-
ceous macrofossils collected by members of the Palaeontol-
ogy Department of The Natural History Museum and others
during their investigations of the Qahlah and Simsima Forma-
tions in the United Arab Emirates-Oman mountain area. The
photograph above shows the spectacular unconformity at an
exposure | km south of Jebel As-Safir in the Jebel Huwayyah
area, between the sub-vertical Jurassic cherts of the Haliw
Formation, and the overlying near-horizontal carbonate
sequence of the late Upper Cretaceous Simsima Formation.
The Haliw Formation is part of the Hawasina Group that
contains mantle-derived ophiolites elsewhere, and is an
obducted nappe complex that was thrust into position during
the mid-Upper Cretaceous and forms the basement beds
below the unconformity. The rudist-rich Upper Campanian/
Lower Maastrichtian Qahlah Formation is absent at this
locality, being overlapped by the Simsima Formation which

Issued 30 November 1995 i

lies directly on the truncated beds of the basement. The |
Simsima Formation yielded most of the other macro #0Say
including 45 species of echinoids belonging to 33 genera,
which is one of the most diverse assemblages of echinoids in
the Upper Cretaceous known anywhere in the world. The
field collecting and work were made possible by a continuing
grant to Peter J Whybrow, The Natural History Museum,
from the Abu Dhabi Company for Onshore Oil Operations,
and in particular we would like to thank Dr Terry Adams, Mr)
David Woodward and Mr Kevin Dunne, successive General
Managers of ADCO for their support and encouragement. In
addition we thank Mr Nabil Zakhour, Head of Public Affairs
and Dr Jose E de Matos, Senior Geologist, ADCO, for their
sustained assistance. The ADCO grant for research in and
around the Emirate of Abu Dhabi forms part of the Natural
History Museum’s ‘Global Change and the Biosphere’
research theme.

Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):91-119

Issued 30 November 1995

Late Cretaceous carbonate platform faunas of
the United Arab Emirates-Oman border

region

A.B. SMITH, N.J. MORRIS and A.S. GALE
Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD,

U.K.
W.J. KENNEDY

Department of Earth Sciences, University of Oxford, Parks Road, Oxford OX1 3PR, U.K.

CONTENTS
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ithotaciecrancbaunalkassem laces a aceedeeeeccete sss ohascce ei sneect ee eener eeaehccoeteeceee icc en eS 95
General palaccenvironmentalisynthesismepestessmererecetes.-< ceeeseeet etcetera cee ee 107
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Synopsis. The stratigraphy and macrofauna of the Qahlah and Simsima Formations is described for 14 sections
along the western margins of the North Oman Mountains between Al Ain and Al Dhaid. These are the earliest
autochthonous deposits following emplacement of the Semail ophiolite complex and are dated, on the basis of
ammonites, inoceramid bivalves and echinoids, as latest Campanian to Maastrichtian in age. Deposition over a
deeply weathered surface of ultrabasic rocks commenced with nearshore conglomerates, grits and cross-bedded
sands which, in places, have an in situ bivalve fauna. Clastic input was abruptly terminated by a sea-level rise and
replaced by carbonate deposition in the early Upper Maastrichtian. The basal carbonate bed is composed of coarse
shoreface reefal debris formed during rapid transgression. This is overlain by a highly fossiliferous series of muddy
carbonate sands deposited in shallow water around wave-base. There is indirect evidence for algal stands from the
associated macrofauna, and level-bottom thickets of corals/rudists are present. Upper beds are well-washed
carbonate sands rich in larger benthic foraminifers but with few macrofossils that were deposited in broad subtidal
flats. In places these are overlain by deeper shelf-basinal marls. Bed by bed faunal lists for each section are provided.

INTRODUCTION

The Oman Mountains form a prominent arcuate range along

the northeastern corner of the Arabian Peninsula. Their

geological history is complex, but work by Glennie et al.

(1974), Glennie (1977), Hughes-Clarke (1988) and contribut-

ers to Robertson et al. (1990) has done much to improve our

understanding of this region. The rocks forming the Oman

Mountains can be divided into seven major geological units,
ranging in age from late Proterozoic to early Tertiary. Of
direct interest here are the Hawasina Group, the Semail
Nappe and the Maastrichtian to early Tertiary autochthonous
marine clastics and carbonates. The Hawasina Group is
formed of tectonic slices of Permian to mid-Cretaceous
sedimentary sequences deposited over the ocean floor and
continental slope of the Arabian continent. They were
dbducted onto the continental margin of the Arabian plat-
orm during the Upper Cretaceous. The Semail Nappe repre-
‘ents a massive slice of former oceanic crust, generated by

) The Natural History Museum, 1995

subduction-related spreading during the Cenomanian-
Turonian and emplaced before the Upper Campanian. Both
are overlain by marine sediments of Maastrichtian to Palae-
ocene age. Initially these beds were deposited around the
newly emergent margins of the Arabian shield, following the
obduction event. Later, a broad carbonate platform formed
over the region as it subsided. Only remnants of this once
widespread succession now remain, forming small jebels
along the western margins of the Oman Mountains.

The late Cretaceous to early Palaeocene rocks are conven-
tionally divided into three units (Skelton et al. 1990), which
are, from bottom to top:

1. The Qahlah Formation — a marine clastic sequence of
sands and conglomerates of late Campanian or early
Maastrichtian age.

2. The Simsima Formation — a platform carbonate sequence
of Maastrichtian age.

3. The Pabdeh Group — a basal limestone conglomerate of
reworked Simsima Formation with an erosive base, of

presumed early Palaeocene age, overlain by thin-bedded
basinal marls of late Palaeocene age.

Until recently little was known of the fauna of the late
Cretaceous rocks of this region. The first published account
of Cretaceous fossils was that of Carter (1852), who described
late Cretaceous sediments of the Hadramaut region of south-
east Oman. Parts of the fauna collected by Carter were
described by Duncan (1865) and are Cenomanian in age.
Lees (1928) gave the first authoritative account of the geology
of the Oman Mountains and provided much new palaeonto-
logical data. He described late Cretaceous faunas from sev-
eral localities, including Jebel Bu Milh (‘Jabal al Milah in
Wadi Sharm’), where he recorded and described 39 taxa of
gastropods (‘the product of an hour’s collecting’). Clegg
(1933) also described a small number of late Cretaceous
species from Oman, but without stratigraphic details.

In the past few years there has been renewed interest in the
geology and palaeontology of the late Cretaceous deposits
around the fringes of the Oman Mountains. The regional
setting of these deposits was outlined by Alsharham & Nairn
(1990, 1994), who also described lithofacies and listed micro-
fauna for type sections. The late Cretaceous stratigraphy and
faunas from the south western Oman Mountains (Dhofar
region, Oman) were studied by Platel & Roger (1989), Roger
et al. (1989) and Roman et al. (1989). Late Cretaceous
ammonites, echinoids, foraminifera and calcareous algae
were described from the central Oman Mountains (Smith er
al. 1990, Kennedy & Simmons 1991). To the north, along the
United Arab Emirates-Oman border region, Skelton er al.

Fig. 1 Map of the Arabian Peninsula showing the area of study.

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

(1990) studied key late Cretaceous sections from both a
sedimentological and palaeontological standpoint. They pub-
lished lithostratigraphic sections and faunal lists for these
sections, with particular emphasis on the rudist bivalve fau-
nas, from which they were able to provide the first detailed
assessment of late Cretaceous palaeoenvironments for the
region, demonstrating that the carbonate succession recorded
a variety of facies ranging from intertidal to shelf basinal
settings. Finally, there are a series of papers documenting
part of the diverse echinoid fauna (Ali 1989, 1992a,b) and a
few of the molluscs and corals from this region (Ghalib, 1989,
1990; Metwally 1992).

Our interest in the faunas of this region began in 1984,
when Dr. S. Nolan (then at the University of Swansea) and
Dr. P.W. Skelton (Open University) brought back their
collection of late Cretaceous fossils for identification. This,
together with material brought to the Natural History
Museum for identification by amateur collectors (notably,
Mrs Valerie Chalmers), alerted us to the importance of the
Maastrichtian faunas of the western fringes of the Oman
Mountains. In April, 1991, A.B. Smith carried out a prelimi-
nary survey of some late Cretaceous fossil localities along the
Oman-United Arab Emirates border. The echinoid fauna
proved to be exceedingly rich and also remarkably well-
preserved, and without doubt represents the most important
Maastrichtian Tethyan echinoid fauna yet known.

During this initial survey it became rapidly apparent that
the molluscan and coral faunas were also exceedingly rich and
likely to be of equal importance, both in terms of new taxa
and their significance for understanding late Cretaceous

CUA

Arabian Peninsula

|
E
|

}
|
t
|

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

benthic community structure. Indeed, the small collections
brought back to London for examination here by relevant
experts generated considerable interest. Consequently a sec-
ond expedition was mounted to study the entire macrofauna
from these important sections and to investigate the lithofa-
cies and palaeoenvironments in more detail. In January, 1992
three of us (ABS, NJM, ASG) spent two weeks exploring,
logging and collecting from the late Cretaceous outcrops that
form the western margins of the Oman Mountains along the
United Arab Emirates Oman borders region. It is the results
of these two bouts of fieldwork that form the subject of the
following papers.

STUDY AREA

Our study area lies along the border between the United
Arab Emirates and the Sultanate of Oman (Fig. 1). Outcrops
in this region comprise small, generally outlying jebels along
the western fringes of the Oman Mountains. Collection and
logging was carried out at eight sites ranging from Jebel
Huwayyah in the south to Jebel Faiyah in the north (Fig. 2).
Sections were measured at each jebel and macrofauna col-
lected or noted bed by bed. Details of the outcrops studied
are as follows:

1. Jebel Huwayyah (Figs 2, 3). Two sections were logged and

' their faunas collected systematically.

Jebel Huwayyah, section 1. Southeastern corner of the

| U-shaped Jebel immediately to the north of the dirt track and
_ about 3 km SE of the northwestern corner of the jebel, 10 km

NE of Al Ain, United Arab Emirates. Map reference:
Buraymi Sheet 1:100,000 NG—40-140; grid reference

| 842,878.

' Jebel Huwayyah, section 2. Western face of the eastern limb
of the jebel approximately 2 km SSE of the northwestern
corner of the jebel, 10 km NE of Al Ain, United Arab

| Emirates. Map Reference: Buraymi Sheet 1:100,000,

_ NG-40-140; grid reference 823,877.
|

2. Jebel Bu Milh (Fig. 3). Two sections were studied in detail.
Jebel Bu Milh, section 1. Western face of a prominent ridge at
| the northwestern tip of the jebel, 10 km NW of the village of
-Mabdah, Oman. Map reference: Buraymi Sheet 1:100,000
NG+0-140; grid reference 895,075.

Jebel Bu Milh, section 2. Southeastern corner of a prominent
knoll, isolated from the main jebel at its northern end, some
10 km NW of Mabdah, Oman. Map Reference: Buraymi
Sheet 1:100,000 NG-40-140; grid reference 906,075.

3. Jebel Rawdah (Figs 2, 4B). Six sections were examined in
detail and a further two briefly investigated but not logged or
systematically sampled. Logged sections are as follows:

lebel Rawdah, section 1. Slope and cliff at the eastern end of
“he northern flank of a valley some 50 m east of the head of
he valley, 3 km east of quarry weigh-bridge and site office,
jebel Rawdah, east of Al Madam, United Arab Emirates.
dumayni sheet NG-40-14A 1:100,000; grid ref. 925,528.

ebel Rawdah, section 2. Slope and cliff on northern side of
‘ebel Rawdah, about 2 km east of quarry weigh-bridge and
ite office, Jebel Rawdah, east of Al Madam, United Arab
‘mirates. Sumayni sheet NG—40-14A 1:100,000; grid ref.

Al Hair

Jepel
Huwayyah

Fig. 2. Locality map showing the position of the four major jebels
studied (asterisked) where there are important outcrops of late
Cretaceous sediments. For regional placement see Fig. 1.

913,544.

Jebel Rawdah, section 3. Slope and cliff on south side of
valley from 200 to 600 m east of the head of the valley, about
3 km east of quarry weigh-bridge and site office, Jebel
Rawdah, east of Al Madam, United Arab Emirates. There
were three measured sections, labelled from west to east a, b,
and c. Sumayni sheet NG—40-14A 1:100,000; grid refs
932,528 (section 3a): 928,527 (section 3b) and 927,527 (sec-
tion 3c).

Jebel Rawdah, section 4. Slope and cliff at the eastern end of
the northern flank of a valley, 600 m east of the head of that
valley, 2.5 km east of quarry weigh-bridge and site office,
Jebel Rawdah, east of Al Madam, United Arab Emirates.
Sumayni sheet NG-40-14A 1:100,000; grid ref. 928,531.

4. Jebel Buhays (Figs 2, 4A). Two sections were logged and
their faunas systematically collected. A third section (2),
exposing the lowest beds of the sequence was impossible to

\4Jebel Qatar }:/4 % : |

Fig. 3 Detailed locality map for Jebel Huwayyah and Jebel Bu Milh. 1, 2 = studied sections. For regional placement see Fig. 2.

log, but collections were made from the scree slope.

Jebel Buhays, section 1. East face of the most northerly hill
forming Jebel Buhays, 4 km north of Al Madam, United
Arab Emirates. Dhayd Sheet 1:100,000, NG—40-107;
780,681. A second section 300 m to the south and forming
another small hill was also examined and collected from, but
no detailed section was made.

Jebel Buhays, section 2. Scree slope at the southwestern
corner of Jebel Buhays, 4 km north of Al Madam, United
Arab Emirates. Dhayd Sheet 1:100,000, NG—40-107; grid
ref. 779,668. No measured section could be made, but the
beds are almost vertical here and the scree material is all
derived from the lowest few metres of the sequence.

Jebel Buhays, section 3. Northeastern corner of Jebel Buhays,
just southeast of a television mast, 4 km north of Al Madam,
United Arab Emirates. Dhayd Sheet 1:100,000, NG—40-107;
grid ref. 788,670.

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

U
U
‘

5. Jebel Thanais (Fig. 4A). Northeastern side of the jebel,
about 4 km north of Al Madam, United Arab Emirates. |
Dhayd Sheet 1:100,000, NG—40-107; grid ref. 783,699.

6. Jebel Aqabah (Fig. 4A). Southwestern face of jebel,
forming a steep cliff about 200 m northeast of the tip of Jebel
Thanais and about 4 km north of Al Madam, United Arab
Emirates. Dhayd Sheet 1:100,000, NG—40-107; grid ref.
785,698. |

7. Jebel Faiyah (Figs 2, 4A). One section was logged in detail.
A further three were explored but found to be unprofitable:
collections were made from only one.

Jebel Faiyah, section 1. Eastern scarp face from the southern,
nose of the jebel northwards for 500 m, 5.5 km north of Al
Madam, United Arab Emirates. Detailed logs were made a
the northern end of the section (section 1a) and approxi

L Jebel Buhays }
aA

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

mately midway along (section 1b). Dhayd Sheet 1:100,000,
NG—~40-107; grid ref. 800,697 to 802,702.

Jebel Faiyah, section 2. Eastern scarp face of the jebel
approximately 3 km NNE of the southern tip of the jebel, and
8 km N of Al Madam, United Arab Emirates. Dhayd Sheet
1:100,000, NG—40-107; grid. ref. 806,722. All but the lowest
2 metres of outcrop was highly indurated and thus fossils
collected come from only these basal beds.

8. Qarn Murrah (Fig. 4A). Northeastern slope of the garn,
some 8 km west of the northern tip of Jebel Faiyah and 15 km
north of Al Madam, United Arab Emirates. Dhayd Sheet
1:100,000, NG—40-107; grid ref. (approx.) 760,795. The
section here was small and the rocks hard and, well-lithified,
making collecting difficult. No section was logged.

LITHOFACIES AND FAUNAL ASSEMBLAGES

A total of 14 sections were logged and their macrofauna
recorded. Lithological samples were collected systematically
from the thickest sequence, (Jebel Rawdah, section 2), and
sectioned for petrographic analysis. The lithological descrip-
tions given for other sections are based solely on field
observations and are consequently of a preliminary nature.
The results of this work are summarized in the measured
sections (Figs 5-11) and in the faunal lists of the Appendix.
As the successions in the various jebels differ significantly
and correlation between jebels was not initially obvious, each
succession is documented in turn before attempting a synthe-
sis.

Qarn Murrah

N
1 ar
wal
Qarn ee
Mulayh tbe a
2S

3, 4 = studied sections. For regional placement see Fig. 2.

Jebel Rawdah (Figs 5-7)

1. Sedimentary lithofacies at section 2. From field logging and
petrographic analysis we recognize the following major litho-
facies:
Facies 1. The succession commences with an ultrabasic clast
conglomerate (bed 1). This can reach more than 10 m in
thickness in places, and comprises well-rounded clasts of
mean size 10-20 cm. There are rare rounded fragments of
rudist and occasional acteonellid gastropod shells. Similar
sediments in the region have been interpreted as beach
deposits by Skelton e¢ al. (1990), a view supported in our
interpretation.
Facies 2. There then follows a mixed clastic-bioclastic coarse
sand, with relatively well-rounded coarse sand to gravel-sized
clasts, ca. 1 m thick (beds 2-4). This includes up to 20%
ultrabasic sand. The high sphericity and excellent sorting of
the clasts indicates these sediments accumulated in a very
high-energy environment; either intertidal or very shallow
subtidal. We interpret these as shoreface sands.
Facies 3/4. The majority of lower beds in the succession are
poorly sorted mollusc-coral-foraminiferal packstones that
contain components of three size classes: (i) clay and silt-
grade carbonate, now replaced by a weak ferroan calcite
microsparite cement; (ii) well-rounded, fine sand-grade par-
ticles, with a sizeable component of ultrabasic grains in the
lower beds, as, well as rolled and bored mollusc and echino-
derm debris; (iii) diverse larger bioclasts which vary in
composition from bed to bed. There are two broad facies
distinguishable in hand specimen on the basis of the major
clast component.

Facies 3 is a poorly sorted calcarenite rich in mollusc clasts
(especially rudist clasts) as well as obvious ultramafic sand-

Detailed locality map for Qarn Murrah and Jebels Buhays, Thanais, Aqabah and Faiyah (map A), and Jebel Rawdah (map B). 1, 2

Figures 5 - 11. Measured sections through the late Cretaceous Qalah and Simsima Formations. Locality
details for each named section are given in Figures 2 - 4 and in the text. Logs are drawn to a scale of

5 mm = 1 m for Figs 5, 6,9 - 11 and 4 mm = 1 m for Figs 7 and 8. Lithological data are based on field
observations except for Jebel Rawdah, section 2. Faunal assemblages noted in the right-hand column
are also largely based on field observations; full faunal lists for each section are given in Appendix 1.

Symbols used are as follows:-

Bioturbation

Poorly-sorted bioclastic
packstones with major

grade clasts. This makes up the lower beds (beds 5-10) and
includes various thin coquina lenses and horizons of coarse
coral and rudist debris.

Facies 4 is characterized by an abundance of orbitoid
foraminiferan clasts, with fragments of the rudist Dictyopty-
chus common in lower beds (beds 11-12) and of corals in
upper beds (beds 15-19). Furthermore, in these higher beds
there are also layered blebs of mud-grade carbonate, incorpo-
rating sand-grade bioclasts, which are storm-reworked pellets
of mud-grade sediment.

Skelton er al. (1990) interpreted facies 3 and 4 as deposits
formed in a point bar and tidal channel system. However, the
original presence of significant quantities of mud-grade car-
bonate (probably algal aragonite) in this facies, together with
its overall poor sorting is taken as evidence of deposition
below normal wave-base. The sand-grade component, which

Siliciclastic sands &
conglomerates

Well-sorted bioclastic

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

Coarse, poorly sorted
bioclastic wackestones

Red-weathering marly
seams and wackestones

orbitoid foraminifer packstones |
content |
Weathered serpentenite Rudists ©o@ Rudists ;
(Semail complex) ANI (Dictyoptychus) ( Hippurites)
Rudists Infaunal bivalves = Bivalves |
(large Vaccinites) © (Scabrotrigonia) ‘S~ (undifferentiated)
Gastropods ae Gastropods Gastropods :
(Campanile) (Acteonella) @ (undifferentiated) |
“, 4 |
Corals CS Stromatoporoids SOF Echinoids
Loftusia (benthic
LL
ASE NCCES foraminiferan) |

is well-rounded and was originally well-sorted, was probably |
washed in from shallower water (intertidal to shallow sub-
tidal) by storm activity.
Metre- to several metre-scale bedding cycles (most obvi-

ously expressed in banded rhodolite development) are appar-
ent in this facies. The origin of such cyclicity is enigmatic, but
the deeper-water setting proposed above militates against
their being tidal channel in origin.

Facies 5. The upper part of the succession (comprising beds
19-28) is composed of well-sorted, well-rounded sand-grade
bioclasts, grain supported and originally containing little or
no calcite mud. There are rare larger (up to 1 cm) bioclasts.
The foraminiferal component is dominant (50-60%) and
includes both broken and rolled orbitoids, entire small rota- |
lines and miliolines. Up to 10% dasycladacean algae also
occurs. This facies was deposited significantly above wave-

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

Jebel Rawdah 2 (lower part)

Massive well-sorted
packstones with
scattered orbitoids

Prominent parting

Massive well-sorted
packstones with
scattered orbitoids

Nodular well-sorted
bioclastic sand
Red marly parting

Nodular weltsorted
bicckstic sand

Bioturbated orbitoid-
rich bioclastic
Packstone
Orbitoid-rich
wackestone

Poorly sorted
bioclastic
wackestones rich in
orbitoids and with
rhodolite bands

Red-weathering, flaggy
bedded silty cakcarenite

Poorly sorted
orbitoline-rich
biociastic
wackestone

Nodular, coarse
bioclastic wackestone
with orbitoids

Orange parting

Grey poorty-
sorted bioclastic
Ppackstone with
siliciclastic grains

Sandy calcarenite

Boulder conglomerate
Sandy calcarenite

Boulder conglomerate

Campanile 28

Occasional Hemipneustes

Echinoid level - Hemipneustes,
Pygurostoma, Petalobrissus, Noetlingaster;
Barbatia, Pycnodonte, Campanile and
gastropod moulds

Pygurostoma, Noetlingaster, Hemipneustes

Petalobrissus, occasional Globator, Faujasia;
Large Astrogyra; Dictyoptychus, Tylostoma
and other molluscs

27
Abundant Cunnolites and gastropods

Loftusia common
Bournonia

Petalobrissus, occasional Hemipneustes;
Loftusia in levels; gastropods common;
Cunnolites moderately common

Hardouinia, Hemipneustes, Petalobrissus

Alternating Dictyoptychus and Cunnolites
beds, with Loftusia and Petalobrissus

Giant Dictyoptychus

Cucullaea, Pholadomya; Petalobrissus, 25
Globator

Noetlingaster, Zuffardia, Petalobrissus;
Cunnilites; Modiolus

Abundant Petalobrissus; occasional mollusc
debris

24
Echinoid-ich level with Porosoma

Bournonia abundant

Lots of broken moliuscan debris, rare
Dictyoptychus and stromatoporoids; rare
Faujasia, Goniopygus and Orthocyphus

Molluscs, Faujasia, rare Goniopygus, 23
Orthocy phus

Echinoid-rich level

Molluscan debris

Echinoid-rich level

Rolled and bored stromatoporoid heads,
ocassional compound corals and
Dictyoptychus

Molluscan debris; abundant Bournonia;
Faujasia, Goniopygus, Porosoma, Salenia

Rudist and other molluscan debris; Faujasia

Jebel Rawdah 2 (upper part)

Massive-bedded
wel-sorted packstone
with scattered
orbitoids and other
bioclastic debris

Nodular, red-weathering
silty calcarentte

Massive well-sorted
packstones with
scattered orbitoids

Fig.5 Measured section made at Jebel Rawdah, section 2 (see Fig. 4B for locality).

Large Dictyoptychus and other
poorly preserved rudists abundant

Occasional levels of echinoid
and molluse debris seen in
cross-section

Petalobrissus, Hemipneustes,
Noetlingaster, Pygurostoma,
Faujasia; Campanile

Noetlingaster

Campanile

base in a high energy environment, as indicated by the high
degree of sorting and lack of mud-grade carbonate.

2. Succession and faunal assemblages (Figs 5-7). Lateral
variation is considerably greater here than at any other jebel.
Of particular note is the marked attenuation of the entire
lower part of the succession eastwards along Jebel Rawdah 3
(Fig. 7). However, although there is marked variation in bed
development from one section to the next, the same general
succession can be identified at them all.

The basal pebble to boulder conglomerate varies tremen-
dously in thickness, being best developed at Jebel Rawdah 3a
and thinnest at Jebel Rawdah 4. There is very little in the way
of sand and grit lenses developed, the entire sequence being
exceedingly coarse. However, rare Acteonella are present in
the upper part of the sequence at Jebel Rawdah 3.

The coarse siliciclastics are succeeded by a thin transitional
sandy calcarenite facies quickly followed by grey, shelly
bioclastic limestones with obvious scattered sand-sized grains
of ophiolite. These basal calcarenites are relatively coarse
and are well-lithified. At several horizons the beds contain
rolled and often bored heads of the compound coral Actina-
cis, some up to 30 cm in diameter, as well as a variety of
allochthonous compound corals and mollusc debris. The
infaunal cassiduloid Faujasia eccentripora is the only obvious
autochthonous element in these beds. The basal bed at Jebel
Rawdah 4 is notable for the abundance of transported
hippuritid and Durania rudists. Within the succession at
section 2 there are two closely-spaced shell-rich partings with
many echinoids, notably Goniopygus, Phymotaxis and
Orthocyphus.

Towards the top of this calcarenitic succession is a promi-
nent, orange-weathering, silt-enriched parting (top of bed 10,
section 2). This bed is thickest at Jebel Rawdah 2 and has the
same echinoid fauna as found below. The overlying beds are
still bioclastic calcareous sands but are more poorly sorted,
and in places include significant amounts of sand-grade
siliciclastics (e.g. section 1, bed 1). They contain the solitary
discoidal coral Cunnolites and the infaunal bivalves Cucul-
laea, Pholadomya and, in places, Scabrotrigonia. Orbitoid
Foraminifera occur but only in minor abundance.

Overlying these poorly sorted bioclastic limestones come
orbitoid-rich packstones, with abundant specimens of the
infaunal cassiduloid echinoid Petalobrissus. At the base there
is a distinctive infaunal echinoid assemblage dominated by
the cassiduloid Zuffardia and the epifaunal regular echinoid
Noetlingaster. One to three metres above the base of
orbitoid-rich limestones there is a prominent horizon of very
large examples of the rudist Dictyoptychus, probably in life
position. The succeeding 2-3 metres at Jebel Rawdah 2
consist of beds with Dictyoptychus alternating with beds rich
in the solitary discoidal coral Cunnolites, the larger benthic
foraminifer Loftusia, and the infaunal cassiduloid echinoid
Petalobrissus. It is at about this level that the ammonite
Brahmaites (Anabrahmaites) vishnu (Forbes, 1846) was
found at Jebel Rawdah 1. A second major silt-enriched level
occurs above this, and these flaggy beds yield a fauna of large
infaunal cassiduloid and holasteroid echinoids (Hardouinia,
Hemipneustes).

There then follows a succession of rather fine, muddy,
thoroughly bioturbated, bioclastic limestones with orbitoid
Foraminifera and bands of rhodolite nodules. There are
occasional infaunal holasteroid echinoids (Hemipneustes),
but the predominant component of the fauna are the solitary

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

discoidal corals (Cunnolites and Asteraea) and a variety of
gastropods, together with the larger benthic foraminifer
Loftusia.

The succeeding bed (bed 19, section 2) contains large
compound corals (Astrogyra) and rudist material. This bio-
clastic limestone also has occasional large infaunal echinoids
and a mixture of other molluscs.

The next few metres are composed of well-sorted, red-
weathering, silt-rich calcarenites. They have a significant fine
siliciclastic content, particularly in the Hemipneustes beds
(bed 21, section 2).

There then follows a thick and fairly monotonous sequence
of well-sorted calcarenites with sparse to moderate numbers
of orbitoid Foraminifera. This facies has little in the way of
macrobenthos, other than the occasional Dictyoptychus,
Noetlingaster or Campanile. There is, however, one distinct
nodular, red-weathering, silt-enriched calcarenite level which
has a diverse fauna of infaunal cassiduloid and holasteroid
echinoids towards the top.

Large, poorly preserved rudists occur in abundance at the
top of the sequence and the succession is terminated by an
unconformity.

3. Palaeoenvironmental interpretation. It is clear that at Jebel
Rawdah local sedimentation patterns were controlled by
topographic variation of the underlying sea-floor. In particu-
lar, the south-eastern corner appears to have been relatively
starved of sediment compared, for example, to the northern
flank of the jebel.

In environmental terms deposition commenced with
pebble conglomerates laid down in a near-shore high-energy
environment around the newly emergent obduction complex.
These were replaced rapidly, as the obduction complex
subsided and the region became flooded, by coarse, poorly
sorted carbonate sands. These contain significant amounts of
reefal debris and represent immediately offshore sands accu-
mulating below wave-base. The following sequence of mixed
orbitoid-rich platform shoals alternating with more protected
platform bioclastic sands with their gastropod and solitary
coral fauna probably represent local variation between topo-
logical highs and dips over a broad carbonate platform in
probably no more than 10-20 m water depth. Towards the
top of the section a regressive phase is marked by a brief
period of patch-reef development and the influx of fine
siliciclastics. The remainder of the succession is composed of
shallow-water calcarenites formed within active wave-base
and supporting only a sparse fauna.

Jebel Huwayyah.

1. Lithological succession and faunal assemblage (Fig. 8). The
general succession is similar throughout the jebel, though
with some lateral variation in bed thicknesses. The lowest
beds, best seen at section 1, consist of poorly sorted silts and
sands with rare lenses of the oyster Acutostrea. The succes-
sion passes up into pebble- and grit-sized conglomerates and
cross-bedded siliciclastic sandstones, the pebbles being pre-
dominantly igneous in origin and well-rounded. Bed 7 is of
particular interest because it contains broken fragments of
thick-shelled rudists, and occasional pebbles that are
encrusted by Acutostrea or, more rarely, by small compound
corals.

After a small gap in exposure there follows a succession of
highly bioturbated, poorly sorted, brown-weathering, silt-rich

Jebel Rawdah 1

More massive-bedded ,
packstones; well-sorted

Nodular boturbated
orbitoxd-rich bwclastic
packstone

Orange-weathering
rubbty parting

Nodular-bedded, poorty-

and rhodolite bands

Reddish parting
Brown-weathering,

grains

with siliciclastics

sorted, coarse bioclastic
packstone with orbitoids

Orbitoid - rich, bioclastic
packstone, poorly sorted,

bioturbated, poorly sorted
packstone with siliciclastic

Poorly sorted packstone

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

Coenholectypus

?Proraster (in cross-section)
Agerostrea, Pycnodonte

Rare Dktyoptychus

Scattered gastropods, occasional
large coral heads, Spondylus

Hattopsis, Hemipneustes, Noetlingaster;
discoidal corals; occasional Loftus and
Dictyoptychus

Pycnodonte, Neithea, Acteonella, large
infaunal bivalves, lots of gastropods
and discoidal corals; Loftusia;
ammonites

Large Dictyoptychus; Globator,
Petalobrissus, Orthopsis

Faujasia-rich level

Abundant Scabrotrigonia, Cucullaea
and Cunnolites

Abundant Pycnodonte

Fig.6 Measured sections made at Jebel Rawdah, sections 1 and 4 (see Fig. 4B for locality).

Jebel Rawdah 4

Prominent red marl
Parting

Prominent red marl
parting

More massive bedded

Hemipneustes

Faujasia, Coenholectypus

Dictyoptychus

Globator, Pygurostoma,
Coenholectypus

bioclastic packstones with

scattered orbitoids

Rubbly, coarse
brown-weathering
sandy wackestone

Rubbly, coarse,
poorly-sorted
packstone

More massive
orbitoid-rich
bioclastic
packstones

Nodular
orbitoid-rich
biockastic packstone

Soft, bioturbated marl
seam

Orbitoid-rich

bioclastic packstone

Hard nodular rhodolite-
rich bed

Red marly calcarenite

Rubbly, poorty-sorted
bioclastic packstone
with orbitoids

Rubbly, poorty-sorted
biociastic packstone
with orbitoids

Red sandy marl
Rubbly, coarse

bioctastic limestone
with siliciclastics

Hard sandy calcarenite
packstone

Orthopsis, Salenia, Globator

Scabrotrigonia, Pycnodonte

Pycnodonte

Scabrotrigonia, Pycnodonte

Globator, Orthopsis

Dictyoptychus

Dictyoptychus, Globator, Hemiaster

Noetlingaster common, Globator,
Petalobrissus, Coenholectypus

Lophids and Campanile
Dictyoptychus
Globator

Phymechinus, Globator, Orthocyphus,
Petalobrissus

Globator, Petalobrissus, Faujasia,
Glyphopneustes
Globator

Orthocyphus, Faujasia, Globator,
Petalobrissus

Bournonia, oysters, gastropods

Abundant rudists (Hippurites,
Dictyoptychus), gastropod and bivalve
debris

100

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

Jebel Rawdah 3c

Jebel Rawdah 3b

Massive, well-sorted orbitoid
Packstone with chert nodules

Jebel Rawdah 3a "1
Massive red-weathering
Packstone with orbitoids

Thin-bedded, red-weathenng,
silt-rich wackestone

Relatively thick-bedded
orbitoid-rich packstone

More massive orbitoid-nch
packstone

Red weathering parting

Nodular, bioturbated, poorly
sorted packstone with
scattered orbitoids and
thodolite bands

Coarse, poorty sorted
packstone

== —_— — —_ —_— Ss
Coarse conglomerate with
rare lenses of sands and
grits

Fig. 7 Measured sections made at Jebel Rawdah, section 3 (see Fig. 4B for locality).

Cucullaea, Tylostoma
Agerostrea, Amphidonte, Barbatia
Acteonella

Mecaster, Actinophyma, Barbatia,
Modiolus, Pycnodonte, Amphidonte

Diptyoptychus, Agerostrea, Pycnodonte
Clavagella

Eligmid bivalves, sparse Dictyoptychus

Loftusia, solitary discoxdal corals,
Pycnodonte; Hemipneustes, Mecaster

Strombid gastropods, Barbatia, mytiloids;

Rolled and bored compound coral heads;
Dictyoptychus and other rudists;
occasional Acteonella, pycnodontids

Occasional Acteonella
fragments

Jebel Huwayyah 1

Agerostrea and Pycnodonte

Fine-grained carbonate
sands/silts with chert
nodules and scattered
orbitoids

Proraster

Thallasinoides burrow systems

Agerostrea and Pycnodonte;
occasional solitary corals

Campanile
Oyster debris
More massive,
orbitoid-rich,
packstone

Bioturbated poorty-sorted
bioclastic packstone with

orbitoids Diverse fauna, including abundant gastropods

including Campanile; Globator Pygurostoma,
Orthopsis;. corals, bivalves

Coarse, poorty sorted
bioclastic packstone

Pebble conglomerate

Silty bioclastic
wackestone with
calcareous nodules

Bioclastic packstone

Scattered Loftusia

Abundant Loftusia; Faujasia;
Vaccinites, Spondylus
solitary discoidal corals; rare
compound corals, ammonites

Silty bioclastic
wackestone

Rubbly bedded sandy
bicclastic wackestone

Abundant Loftusia

Pebble conglomerate

Cross-bedded, coarse
sand with strings of grit

Rudist fragments; some pebbles oyster encrusted
(Acutostrea); bryozoans

Pebble conglomerate

Cross-bedded, coarse
sand with strings of grit

Poorly sorted
conglomerates

with cross-bedded sand
lenses

Fine carbonate sand

Sparse Acutostrea
with scattered clasts

Poorly sorted
conglomerates
with sand lenses

Small Acutostrea and Gastrochondrites

Sand with scattered
clasts

le. 8 Measured sections made at Jebel Huwayyah, sections 1 and 2 (see Fig. 3 for locality).
|
|

Jebel Huwayyah 2

Bioclastic packstone
Rubbly bedded sandy
bioclastic wackestone

Rubbly bedded sandy
bioclastic wackestone

Compound corals; Dictyoptychus and

Vaccinites; rare ammonites
Abundant Loftusia and discoidal

solitary corals; diverse molluscs;

Hemiaster

Abundant Loftusia and discoidal
solitary corals;
diverse mollusc fauna

Abundant Loftusia and discoidal
solitary corals;

diverse mollusc fauna;
Coptodiscus, Glyphopneustes

101

102

bioclastic limestones with significant amounts of dark-green,
sand-sized, igneous grains. These beds contain abundant
specimens of the benthic foraminifer Loftusia as well as
scattered molluscan debris and occasional infaunal spatan-
goid echinoids. The large solitary button coral Cunnolites is
also abundant. Within this facies are two prominent marker
limestones that are thicker and better developed on the
western limb of the jebel. They contain a rich fauna of
compound corals, the hippuritid rudist Vaccanites and the
occasional large, tall-spired gastropod Campanile. None of
the rudists are found in life position and the beds appear to
represent allochthonous accumulations. However, the base of
the upper of these beds is typically rich in Spondylus and
Plicatula that are clearly not transported, as most are pre-
served with both valves connected. Ammonites, some quite
large, are occasionally found at this level, presumably trans-
ported into this environment.

Above the Loftusia-rich beds comes a sequence of less
fossiliferous silty bioclastic carbonate sands, which are in turn
succeeded by a thick pebble conglomerate. This conglomer-
ate is composed of very well-rounded clasts, and represents
such an anomalous lithofacies change that it must have
formed through exposure and reworking of earlier conglom-
eratic beds. Above the conglomerate siliciclastics abruptly
disappear and are replaced by relatively clean bioclastic
limestones of the Simsima Formation. This starts with an
initial shell-rich bioclastic limestone, followed by 2 to 3
metres of highly bioturbated and poorly sorted bioclastic
limestones with sparse sand-grade siliciclastics. These sedi-
ments are very fossiliferous, the fauna dominated by shallow
infaunal cassiduloids and holectypoids (Pygurostoma and
Globator), infaunal naticids and other gastropods, including
Campanile. Occasional pectinid bivalves are also present.

Higher beds pass into less fossiliferous and much more
massive and well-sorted, orbitoid-rich carbonate sands and
silts. There is at least one level of thallasinoid burrows. Rare
Dictyoptychus are found here. The orbitoid content of the
sediments decreases upwards so that higher beds, which
contain the epifaunal bivalves Agerostrea, Pycnodonte and
the infaunal spatangoid Proraster, are finer-grained carbon-
ate sands or silts with relatively few orbitoid clasts.

2. Palaeoenvironmental interpretation. The succession com-
mences with unconsolidated subtidal sand and cobble beds
deposited around the margins of the newly emergent obduc-
tion complex. The sequence shallows upwards into the shore-
face facies of bed 7, with increasing cross-bedding and coarser
clastic content. The clastic supply then sharply diminishes,
presumably marking the final flooding of nearby ophiolite
islands, and the initiation of carbonate platform develop-
ment. The poorly sorted Loftusia-rich muddy carbonate
sands represent extremely shallow-water, back bar or
lagoonal deposits formed in a protected environment below
wave-base. Coral-hippuritid thickets were able to develop
locally and occasional open-water ammonites were washed
in. The thin, well-sorted calcarenitic sands with Faujasia may
represent protected beach-face sands.

A brief regression caused by local uplift led to the exposure
of Qahlah Formation conglomerates, or their source rock,
and pebble conglomerates were once more briefly deposited
in a high-energy shallow marine environment (bed 13). This
was short-lived and carbonate platform conditions returned
once again, bringing about the more or less complete elimina-
tion of siliciclastic components. This presumably marks a

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

renewed phase of subsidence with transgressive seas once
more flooding the region and cutting off siliciclastic input.
The initial coarse bioclastic fossiliferous sands (bed 14), we
interpret as deposits formed at or immediately below normal
wave-base. They are replaced immediately upwards by thick
beds of shallower-water orbitoid-rich carbonate sands with
little macrobenthos, formed within wave-base. These are
succeeded in turn by fine carbonate sands and silts suggestive
of shelf-basinal conditions.

Jebel Bu Milh

1. Lithological succession and faunal assemblages (Fig. 9).
The succession begins, as in Jebel Huwayyah and elsewhere,
with a rather thick series of well-rounded pebble conglomer-
ates, grits and coarse clastic sands, with some small-scale
cross-bedding. Although largely unfossiliferous, occasional
clumps of the rudist Durania can be found, apparently in situ.
Towards the top is a 1-2 m thick conglomeratic sequence,
packed with well-sorted shells of the gastropod Acteonella,
showing good current alignment of individual layers. Occa-
sional specimens of the rudist Pseudosabinia also occur. The
succeeding clastic sands are strongly cross-bedded and con-
tain occasional Acteonella specimens, as well as large logs
bored by lithophagid bivalves.

There is then a sharp reduction in the siliciclastic content of
the sediments and the succeeding 1.0-1.5 metres are com-
posed of highly fossiliferous calcarenitic limestones with
sparse sand-sized quartz grains. The basal bed is a fine, silty
calcarenite that is highly nodular and extensively bioturbated.
It contains rare nautiloids and ammonites and a rich gastro-
pod fauna, with naticids and neogastropods dominating. An
erect, branching sponge is a common element of this fauna,
though none appear to be preserved in situ. Above this basal
carbonate bed is a thin, red-weathering, silt-enriched parting,
followed by a harder, less bioturbated shelly bioclastic lime-
stone that contains at its base the large tall-spired gastropod
Campanile in some abundance, together with Acteonella,
Dictyoptychus and other large molluscs.

Succeeding beds are well-sorted orbitoid-rich bioclastic
limestones. These contain little in the way of macrofauna
except for a very distinctive horizon of large hippuritid rudists
in situ, some 4-5 m above the base of the limestone succes-
sion.

2. Palaeoenvironmental interpretation. The succession at
Jebel Bu Milh commences with quartz sands, gravels and
conglomerates representing open-water, nearshore deposits
around the newly emergent obduction complex. The succes-
sion shallows upwards, with the Acteonella conglomerates
representing open shore-face, shell-lag accumulations at wave
base, and the overlying cross-bedded sands with beached |
driftwood representing tidal sandbars. The clastic succession |
is abruptly replaced by carbonate platform deposits, presum-
ably as nearby islands became submerged. A thin deposit of
silty bioclastic limestone rich in naticid gastropods represents
shallow subtidal sands, possibly formed at wave base in a_
protected back-bar environment. The presence of the large
neritid Lisocheilus indicates that shallow rocky substrates lay
in the vicinity. The remaining carbonate succession, by
comparison with beds at Jebel Rawdah, represent shallow
subtidal carbonate sand flats formed above wave base.

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

Jebel Bu Milh 1

Well-sorted orbitoid-
rich bioclastic
packstone

Rubbly level

Orbitoid-rich bioclastic
packstone

Bioclastic packstone

Sands with conglomerate
and grit levels, occasional
cross-bedding

Shel-rich, occasional coral

Scattered molluscan debris

Dictyoptychus and Campanile
Gastropods in cross-section
Abundant Campanile and Tylostoma,

rare Deltonautilus

Abundant Dictyoptychus and
Acteonella; rare Campanite and Durania

Clumps of Durania

103

Jebel Bu Milh 2

Massive-bedded, well-

sorted biockastic

packstone with orbitoids

Welksorted bioclastic Giant Vaccinites

Packstone with orbitoids

Bioclastic orbitoid- Sparse molluscan debris

rich packstone

Shelly bioclastic Campanile, Dictyoptychus, Acteonella

packstone and other molluscs

Nodular poorly-sorted iAbundantinaticd and thiid t d

sandy wackestone lant naticid and cerithiid gastropods,

Orange sand occasional Deltonautilus, sponges

Strongly cross-bedded Occasional bored logs, some large;
sands with grit partings Acteonella also occurs

Cross-bedded
coarse
sands and gravels

Acteonella abundant, forming shell
beds, also some cerithiid gastropods
and the rudist Colveraia

Poorly sorted, cross-
bedded sands and grits
with conglomerate
lenses

Pebble conglomerate

fig. 9 Measured sections made at Jebel Bu Milh, sections 1 and 2 (see Fig. 3 for locality).

104
Jebels Buhays, Thanais and Aqabah

1. Lithological succession and faunal assemblages (Fig. 10).
These three jebels show virtually the same succession and can
therefore be treated together. The initial siliciclastic succes-
sion of pebble conglomerates, sands and gravels is thinner
than at Jebel Rawdah or Jebel Bu Milh. At Jebel Aqabah the
succession commences with a 50 cm well-sorted calcarenite
resting directly on top of slightly weathered ophiolite. This
basal bed is notable for the small, uniform size of its fossils; a
small heteromorph ammonite, Glyptoxoceras sp., was recov-
ered from here. Elsewhere the basal contact is not seen.
Towards the top of the clastic sequence, at Jebel Buhays,
section 1, come 3 to 4 metres of laminar and cross-bedded
sands. These contain a fauna of infaunal venerid bivalves and
small turreted gastropods.

As elsewhere, there is a sudden elimination of siliciclastics
coincident with the onset of biocalcarenite deposition, mark-
ing the base of the Simsima Formation. The limestone
succession begins with a hard, 80 cm thick, sandy bioclastic
shell bed full of the gastropods Acteonella and Campanile, the
echinoid Goniopygus and rudist fragments (Durania and
hippuritid), as well as other molluscan debris. Occasional
compound corals occur at this level, but apparently not in
situ.

The succeeding 3 to 4 metres consist of highly bioturbated,
poorly sorted coarse- to medium-grained bioclastic lime-
stones, with scattered orbitoid foraminifera that increase in
abundance upwards. These beds are rich in infaunal echi-
noids (cassiduloids, especially Pygurostoma, and the holecty-
poid Globator) as well as small epifaunal regular echinoids.
They are also rich in gastropods and bivalves, both infaunal
and soft-bottom epifaunal forms. The only rudist to occur
here in abundance is the small recumbent Glabrobournonia.
Within this succession there is a major red-stained, silt-
enriched layer that immediately overlies a thin bed of fine,
?dolomitized limestone.

There is then a second major shell-lag deposit, full of large
sponge-bored shells of Acteonella and other molluscs. Above
this the bedding becomes much more massive and the lime-
stones better sorted and cemented. Immediately overlying
the Acteonella shell bed is an orbitoid-rich packstone with
rhodolite bands and a low diversity fauna of the rudist
Dictyoptychus and the gastropod Campanile.

A major red-weathering silt-enriched horizon occurs
towards the top of the sequence and is immediately overlain
by a one metre thick brown-weathering silty limestone, rich
in tall cylindriform hippuritids. Associated here is the echi-
noid Codiopsis, indicative of rocky shore habitats. The silt-
enriched bed records a marked increase in the clastic
component at this level. It is overlain by massive-bedded,
well-sorted carbonate sands/silts with orbitoids.

2. Palaeoenvironmental interpretation. The palaeoenviron-
mental setting is similar to that seen elsewhere, with shallow
marine sands, gravels and conglomerates deposited above
active wave base. The basal calcarenite at Jebel Aqabah
probably represents a shoreface sand, as suggested by the
well-sorted clasts and fossils.

These sands, gravels and conglomerates are replaced
abruptly by coarse, poorly sorted bioclastic calcarenites con-
taining a diverse molluscan and echinoid fauna. These beds
are highly bioturbated and were deposited in a protected
environment below wave-base, possibly in a shallow lagoonal

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

setting. The occasional large compound coral near the base
suggests that reefal patches developed nearby. Periodic
minor adjustments in sea-level resulted in red-weathering,
silt-enriched partings and beds, marking temporary influxes
of fine iron-rich silt-grade clasts while thin dolomitic levels
may signify temporary supratidal exposure.

A subsequent regressive phase led to the deposition at
wave base of the Acteonella shell-lag of bed 11 followed by
orbitoid-rich carbonate sands with a low diversity macro-
fauna. We interpret these well-sorted and well-lithified beds
as shallow water platform shoals formed above active wave
base. There was a brief event that created the upper iron-rich
siltstone band (bed 14) which was followed by a bed with
hippurited thickets and the shallow-water echinoid Codiopsis,
indicative of nearshore conditions. Shallow-water carbonate
shoals form the remainder of the succession.

Jebel Faiyah

1. Lithological succession and faunal assemblages (Fig. 11).
Here the sequence overlying weathered ophiolite begins with
pebble conglomerates, with interspersed grit and sand lenses
becoming more common towards the top. Occasional frag-
ments of thick-shelled mollusc are present indicating a marine
origin for the unit.

As in other sections, there is an abrupt change to carbonate

sedimentation, commencing with a 40-50 cm shell bed full of |
Acteonella, together with lesser amounts of rudist fragments —

and other thick-shelled molluscan material. The succeeding 3

metres consist of highly bioturbated, poorly-sorted bioclastic _

limestones, with rhodolites and orbitoid foraminifera, a
diverse fauna of gastropods, small regular echinoids and
Dictyoptychus bands.

There is a major, 10 cm thick, red-coloured, silt-enriched

bed which, as at Jebel Buhays, is interpreted as marking a —
minor change in sea-level with increased erosion of lateritic |

soils from the hinterland. It is succeeded by in situ coral
thickets with associated hippuritid rudists (bed 6, section 1a).
The majority of in situ colonies are erect, branching forms

some 50 cm in diameter, with intermixed erect colonies of |

cylindrical hippuritids. A variety of other colonial corals and
rudists also occur. There is also a moderately diverse fauna of
small regular echinoids, such as Glyphopneustes, and small
gastropods at this level.

Next come a few metres of thick-bedded, orbitoid-rich |

:
|
|
|

ee ee es ee

bioclastic limestones with occasional levels of Dictyoptychus
and rhodolites but little else. There is then a return to |
red-weathering, poorly sorted bioclastic limestones with |
significant silt-grade component (beds 8-10). These beds
appear to be relatively iron-rich. The fauna is diverse and
dominated by the epifaunal bivalves, Pycnodonte and
Agerostrea. Small discoidal corals (Cunnolites), occasional
compound corals, the foraminifer Loftusia and rudist Glabro-
bournonia all occur at this level.

The succeeding thick sequence of rather massive, well
lithified and well-sorted carbonate sands, rich in orbitoid]
foraminifera, but with no observable macrofauna, is tun

cated by an erosion surface.

2. Palaeoenvironmental interpretation. The succession begins
with shallow-water sands and gravels deposited above active
wave base. In places they may even become intertidal or
supratidal (Skelton et al. (1990) reported the presence 0
sedimentary structures indicative of intertidal or supratida

f
4

vl)

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

Jebel
Buhays 1

Massive-bedded
orbitoid packstone

Brown-weathering, silty
fine calcarenite

Red silty wackestone

Massive-bedded, well-
sorted orbitoid
packstone

with rhodolite bands

Shelly cakcarenite

Rubbly bedded,
poorly sorted,
orbitoid-rich
bioclastic
wackestone with
rhodolite bands

?dolomitized Ist

Rubbly orbitoid-
rich wackestone

Red silty wackestone

Orange, bioturbated
poorty-sorted bioclastic
wackestone with some
orbitoids and
rhodolites

Sandy bioclastic
packstone

Pebble conglomerate

Calcareous sand with
grit and pebble lenses

Pebble conglomerate

le. 10 Measured sections made at Jebel Buhays,

Codiopsis; occasional
Dictyoptychus

Small hippuritids common;
Globator rare
Hemiaster

Dictyoptychus

Scattered Dictyoptychus

Large Dictyoptychus, Campanile,
Tylostoma and Neithea

Acteonella shell lag with Neithea,
Bournonia, lophids and Dictyoptychus

Acteonella (small) and other
molluscs; occasional Astrogyra;
Globator common.

Campanile, Plagiostoma, Neithea,
Scabrotrigonia, rare Dictyoptychus,
vermiform gastropod; Phymosoma,
Globator

Small Acteonellids and other
gastropods; Pycnodonte, Bournona;
Cunnolites; regular echinoids,
Globator; decapod claws

Lyria, strombid gastropods, eligmids,
Granocardium, Scabrotrigonia
Campanile, Acteonella, Crassatella,
Pycnodonte, rare Durania fragments;
Porosoma

Mesocallista and small gastropods
in lenses

Jebel Aqabah

Sandy packstone

Coarse sand/gravel
with conglomeratic
lenses

Boulder conglomerate

Well-sorted shelly
calcarenite packstone

sections 1 and Jebel Aqabah (see Fig. 4A for locality).

105

Acteonella common with much rudist
debris and other molluscan material

Infaunal venerid bivalves common,
with low diversity gastropod fauna

Small shelly fossils including Faujasia
and Petalobrissus: Nostoceras

Fig. 11

Jebel Faiyah la

Thick-bedded welt

sorted packstones with
scattered orbitoids and
bands of chert nodules

Bioturbated, silty, poorly-
sorted wackestone; red-
weathering

{ Rhodolite band

Poorly sorted bioclastic
wackestone

Thick-bedded, well-sorted
orbitoid-rich packstone
with rhodolite bands

Well-sorted, thick-bedded
bioclastic packstone with
abundant leptorbitolines

Highly bioturbated
red mart-rich wackestone

Orange poorly-sorted
bicclastic packstone with
scattered orbitoids and
some siliciclastic grains

Well cemented sandy
bioclastic packstone with
rhodolites

Sands and pebble
conglomerates

Pycnodonte, Spondylus, Agerostrea,
Amphidote, Dictyoptychus, Cunnolites,

Noetlingaster, Loftusia

Pycnodonte uncinella

Rare Dictyoptychus

Large Dictyoptychus

Noetlingaster

Branching coral thickets with
hippuritid colonies.

Dictyoptychus

Dictyoptychus, small
gastropods, Hattopsis

Acteonella, small gastropods,
Sparse rudists, turrilitids and
Callianasid claws

Jebel Faiyah 1b

Bioclastic packstone with
orbitoids

Red silty wackestone

Orbitoid-rich packstone with
occasional rhodolite bands

Bioclastic packstone with
orbitoids

Poorly sorted bioclastic
packstone

Reddish marl parting
Fine carbonate sand

Coarse poorly-sorted
bioclastic packstone
Shelly bioclastic packstone

Pebble conglomerate
with sand lenses

Measured sections made at Jebel Faiyah, sections la and 1b (see Fig. 4A for locality).

Loftusia, Pycnodonte, bivalves and
gastropods; Mimiosalenia, Globator

Occasional molluscan debris

Dictyoptychus

Occasional Dictyoptychus

Branching coral thickets in situ;
Hippurites colonies many
gastropods; rare Noetlingaster

Abundant Dictyoptychus and many

gastropods and small echinoids

Acteonella abundant at base

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

environments from these beds). Clastic input ceased sud-
denly, marking a major transgressive event, and was replaced
by carbonate deposition. The carbonate succession began
with a transgressive shell-lag deposit rich in acteonellids, after
which shallow lagoonal conditions were established for a
short period. There was then a minor shift in sea-level,
creating a silt-enriched band, which was immediately fol-
lowed by the establishment of coral-rudist level bottom reefal
thickets and peri-reefal bioclastic sands. These in turn were
quickly replaced by shallower-water bioclastic carbonate
sands with abundant orbitoid foraminifera. These we inter-
pret as shallow-water platform shoals formed above active
wave base. Later, a second transgressive phase, marked by an
input of siliciclastic material, brought a brief return to deeper
water conditions, below active wave base, and the establis-
ment of a more diverse fauna once again. This, however, was
short-lived and shallow-water shoal conditions quickly
returned with the deposition of massive bedded carbonate
sands and the virtual disappearance of benthic fauna.

GENERAL PALAEOENVIRONMENTAL
SYNTHESIS

The autochthonous late Cretaceous succession was deposited
over a deeply weathered surface of ultrabasic rocks, which
must have been exposed subaerially for some time. Initial
deposits were locally derived conglomerates and grits formed
around the shores of the newly uplifted ophiolite massifs.
Pebbles and boulders are well-rounded. Although rudist and
acteonellid debris does form a component of these beds, they
_ probably represent shell coquinas washed onshore from sub-
tidal sand-flats rather than autochthonous fauna. There is
evidence locally for more sheltered shoreface facies with
infaunal bivalves, or for more stable cobble-bottom develop-
‘ment, with an in situ fauna of encrusting oysters and/or
corals. In places, as Skelton et al. (1990) note, the rudist
Durania can be found living in situ in cross-bedded sands.
_ A marine transgression largely drowned these ophiolite
islands and led to the onset of carbonate deposition. During
this transgression, shoreface reefal debris, with mixed hippu-
ritid and radiolitid rudists and massive compound corals, was
deposited as a coarse lag at the base of the transgression. The
‘corals are, for the most part, encrusting forms and are
accompanied by a shallow intertidal to immediately subtidal
tegular echinoid fauna composed of species adapted for life
on hard substrata within the zone of active wave surge.
Acteonellid gastropods are a characteristic feature of such
Shell-lag deposits. Faujasia is the only common infaunal
echinoid in this environment and probably lived in nearshore
or shore-face clean, well-washed sediments. Slightly more
protected sand beds were colonized by non-siphonate infau-
tal bivalves such as Scabrotrigonia and Cucullaea.

This facies was succeeded, as sea-level rose, by a thick
succession of rather muddy sands formed at or below active
wave base. At some levels the large semi-recumbent rudist
Dictyoptychus is common, together with bands of rhodolites,
lense orbitoid foraminiferal aggregates and abundant small
nfauna!l cassiduloid echinoids (Petalobrissus and Zuffardia).
The sea bottom must have been loose and unconsolidated,
ying at or just below active wave base. It regularly received
torm-washed bioclastic material.

107

Elsewhere, solitary discoidal corals and the larger benthic
foraminifer Loftusia occur in profusion together with pycn-
odont oysters and a moderately diverse gastropod assemblage
in what appears to have been a slightly muddier sand.
Infaunal echinoids are more or less absent from this facies,
though the small epifaunal regular echinoid Hattopsis is often
locally abundant and may indicate the presence of algal
stands. The molluscan fauna is dominated by gastropods
(especially the ?algivore Acteonella borneensis and the filter-
feeding Umboniidae gen. nov.) and the epifaunal ‘Pycn-
odonte’ uncinella. We interpret these beds as stable, possibly
algal-bound, sands lying below wave-base. Where this facies
occurs close to reefal thickets there is a much higher diversity
of regular echinoids accompanied by the holectypoid Globa-
tor.

Unconsolidated calcarenite shoals, deposited above active
wave base, form a major part of the upper succession and
contain the lowest faunal diversity. The absence of any
large-scale cross-bedding suggests these formed as shallow,
broad, subtidal expanses of sand. Rotaline and milioline
foraminifera are the predominant bioclasts in this facies,
along with dasycladacean algae. It is in this environment that
the large selective deposit feeding irregular echinoids Hemip-
neustes, Pygurostoma and Stigmatopygus are found. The only
regular echinoid found to inhabit this environment was
Noetlingaster.

At Jebel Huwayyah, however, the highest beds appear
rather finer grained and contain the infaunal spatangoid
echinoid Proraster. These may be local basinal sediments
deposited below wave base, and possibly represent the deep-
est water sediments present in the sequence.

The palaeoenvironmental reconstruction that emerges
from our combined sedimentological and faunal evidence
differs somewhat from that given by Skelton ef al. (1990).
They interpreted the lower part of the Simsima Formation as
tidal deposits, whereas we believe that this is inconsistent
with both the sedimentological and faunal evidence. The
petrography of the sediments indicates deposition below
wave base. This is supported by the diverse echinoid fauna,
which is also at variance with a tidal environment.

The faunal associations that we recognize represent more-
or-less contemporary assemblages that replace one another in
vertical succession as a result of shifting sedimentological
facies. We found very little stratigraphic variation within
single lineages, except in species of the echinoids Hattopsis
and Hemipneustes. Consequently, we believe that the differ-
ences in the assemblages reflect variations in palaeoenviron-
mental conditions over the carbonate platform and represent
coexisting communities.

BIOSTRATIGRAPHICAL RESULTS

Maastrichtian biostratigraphic zonation. The Qahlah Forma-
tion has been regarded as Campanian or early Maastrichtian
in age, and the Simsima Formation as Maastrichtian (e.g.
Skelton et al. 1990). The boundary between Campanian and
Maastrichtian stages has been defined on the basis of a range
of biostratigraphic criteria, involving belemnites, ammonites,
planktonic foraminifera and coccoliths (amongst other
groups). However, recent reviews (Burnett ef al. 1992,
Kennedy et al. 1992, Hancock et al. 1992) have demonstrated

108

that these various data do not correspond. Indeed, Obradov-
ich (1993) concluded that the interval between the widely
adopted planktonic foraminiferal boundary datum (the
extinction point of Globotruncanita calcarata) and the widely
adopted belemnite definition (appearance of Belemnella lan-
ceolata) is at least three million years. There is as yet no
agreed definition of the boundary, and subdivisions of the
stage into Lower/Upper or Lower/Middle/Upper vary widely
between authors. Indeed, in the absence of a clear statement
of which definitions of stage and substage limits are used,
these terms are near meaningless.

Burnett et al. (1992) were able to place the putative
markers for the base of the Maastrichtian in sequence, as well
as certain other key fossil occurrences. Their scheme is as

follows:

(Youngest)

— last occurrence of the nannofossil Quadrum trifidum [or
Tranolithus phacelosus (= orionatus)]|.

— first occurrence of the ammonites Pachydiscus (P.) neu-
bergicus and Acanthoscaphites tridens.

— last occurrence of the nannofossil Broinsonia parca.

— first occurrence of the ammonite Hoploscaphites constric-
tus s.l.

— first occurrence of the belemnite Belemnella (B.) lan-
ceolata.

— last occurrence of the ammonite Nostoceras (N.) hyatti.

— last occurrence of the nannofossil Reinhardtites anthopho-
rus (or Eifféllithus eximus).

— first occurrence of the ammonite Nostoceras (N.) hyatti.

— last occurrence of the planktonic foraminifer Globotrun-
canita calcarata.

— first occurrence of the planktonic foraminifer Globotrun-
cana falsostuarti.

— first occurrence of the nannofossil Reinhardtites levis.

(oldest)

As they note, the first occurrence of the belemnite
Belemnella lanceolata is widely taken to indicate the base of
the Maastrichtian stage, and this is the definition adopted in
the present work. Birkelund ef al. (1984), in their review of
the conclusions of the 1983 Copenhagen Meeting on Creta-

SUBSTAGE

Belemnella casimirovensis

Upper
Maastrichtian

Belemnitella junior

Belemnella occidentalis

Lower

Maastrichtian Belemnella obtusa

Belemnella lanceolata

correlation.

BOREAL BELEMNITE ZONATION

Belemnella fastigata
Belemnella cimbrica

Belemnella sumensis

Belemnella pseudobtusa
Belemnella lanceolata

Fig. 12 Biostratigraphic zonation schemes for the Maastrichtian based on belemnites, ammonites and inoceramid bivalves, and their

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

ceous stage boundaries, noted that it was ‘widely accepted to
keep the base of the Maastrichtian close to the appearance of
Belemnella lanceolata, as this datum is so well-defined and
widely accepted in the Boreal realm. However, there is a
strong need for finer correlation of this boundary level with
the succession in the Tethyan Realm possibly by planktonic
foraminifera or coccoliths’.

Since the work of Burnett et a/. and the partial achievement
of the needs expressed at the Copenhagen Meeting (see also
Schonfeld & Burnett 1991, Kennedy et al. 1992, Hancock et
al. 1992, Hancock and Kennedy 1992) further refinements in
correlation of the base of the Maastrichtian in the classic
Boreal sense with Tethyan sequences have been achieved
through the work of Burnett, Kennedy & Ward (1992),
Hancock et al. (1993), Hancock & Kennedy (1992) and Ward
& Kennedy (1993). These results are summarized in Figure
12. Ward & Kennedy (1993) recognized a threefold ammo-
nite zonation of the Maastrichtian, with a fourth zone of
Pseudokossmaticeras tercense that was possibly in part Cam-
panian, in part Maastrichtian. Taking the base of the Maas-
trichtian at the first appearance of Belemnella lanceolata, the
work of Hancock & Kennedy (1992) and Hancock et al.
(1992) demonstrate that Pachydiscus (P.) epiplectus, Hoplos-
caphites constrictus and P. (P.) neubergicus first occur within
a very narrow interval, such that the base of the Boreal
lanceolata Zone and Tethyan epiplectus Zone are coeval,
within the current limits of biostratigraphic correlation.

For subdivisions of the Maastrichtian we use Lower and
Upper Substages, as is widely accepted by workers in the
Boreal Realm, the base of the Upper Maastrichtian lying at
the base of the Belemnitella junior Zone (Fig. 12).

Anapachydiscus fresvillensis (index species of the second
zone of the Maastrichtian of Ward & Kennedy 1993) first
occurs in the junior Zone in the Netherlands (Kennedy 1987).
The base of the Boreal junior Zone and Tethyan fresvillensis
Zone are coeval, within current limits of biostratigraphic
correlation. Anapachydiscus terminus, index of the highest
Tethyan Maastrichtian ammonite zone, is now known from
the Boreal casimirovensis Zone (as Anapachydiscus aff. fres-
villensis of Birkelund 1993 in Denmark, and, subsequently, in
the Netherlands (Jagt, in press)). For a more refined division

AMMONITE ZONATION
Fi Anapachydiscus terminus |

Anapachydiscus
fresvillensis

Trochoceramus
morgani

Trochoceramus
lanjonaensis

Pachydiscus (P.)
epiplectus

Trochoceramus
monticulae/radiosus

INOCERAMID ZONATION

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

of the Lower Maastrichtian reference will be made to the
zonation of Schulz (1979: fig. 12).

Good palaeontological dating and correlation between the
isolated jebels is very limited. Within each jebel there is
evidence of sedimentation having taken place on an uneven
sea-floor and relativily rapid changes of thickness and facies
are sometimes visible by casual inspection. Larger benthonic
foraminifera are often abundant but apparently controlled in
their distribution by palaeoenvironment. Further careful col-
lecting would be necessary before their stratigraphical poten-
tial is realized. Therefore, here we use evidence from the
ammonite, inoceramid bivalve and echinoid faunas, together
with inferred sea-level changes.

Ammonite biostratigraphy. The small number of late Creta-
ceous ammonites from the United Arab Emirates-Oman
borders region (Kennedy, this volume) provide us with some
constraints on the age of the succession. More weight has
been placed on the pachydiscids than the heteromorphs,
largely because they are better preserved.

The sections at Jebel Huwayyah include at least three
ammonite horizons. The first and lowest contains an uniden-
tified species of Pachydiscus, which was discovered at the top
of the sand and conglomerate sequence of the Qahlah Forma-
tion, just below the Loftusia-rich beds, and only indicates a
late Upper Cretaceous age.

The Loftusia-rich beds themselves have yielded the great-
est number of ammonites, including Pachydiscus (Pachydis-
cus) dossantoi (Maury, 1930), Lewyites ambindense
(Collignon, 1971), Nostoceras (Nostoceras) major Kennedy &
Cobban, 1993, Nostoceras (Nostoceras) sp., and a Libyco-
ceras sp. (not described; photograph only seen). The implied
ages of these ammonites are somewhat contradictory. Pachy-
discus (P.) dossantoi (Maury, 1930) is imprecisely dated
within the Maastrichtian but occurs in the highest of three
ammonite horizons in the Nkporo Shale of south-eastern
Nigeria (Zaborski, 1985). At that locality it is closely associ-
ated with the inoceramids Trochoceramus ianjonaensis (Sor-
nay) and Endocostea coxi (Reyment), which indicate a late
Lower Maastrichtian to early Upper Maastrichtian age.
Lewyites ambindense (Collignon, 1971) is known only from
the lower Upper Maastrichtian, fresvillensis Zone. Nostoceras

| (Nostoceras) major Kennedy & Cobban, 1993 is Upper, but

not uppermost, Maastrichtian, with an inferred fresvillensis
Zone age. The presence of Lewyites ambindense and Nosto-
ceras (Nostoceras) major imply a lower Upper Maastrichtian,
fresvillensis Zone age. However, there are also two species
that suggest an earlier age for at least part of the unit;
Nostoceras (Nostoceras) sp. is uppermost Campanian or
lower Lower Maastrichtian in age, and a specimen of Libyco-
ceras, now in the Arab Emirates University Museum, has a
bifid outer saddle indicative of the earliest of the three
Libycoceras horizons described by Zaborski (1982) from
Nigeria, and a late Campanian age. The age of the Loftusia-
beds could therefore be as old as late Campanian or as young
as early Upper Maastrichtian.

In the northwestern part of the outcrop at Jebel
Huwayyah, close to where it is cut by the road, the lower part
of the Simsima Formation yielded Pachydiscus (Pachydiscus)
_neubergicus neubergicus (Hauer, 1858), and Lewyites ambin-
| dense (Collignon, 1971). Pachydiscus (P.) neubergicus neu-
| bergicus (Hauer, 1858) implies a lower Lower to lower Upper

Maastrichtian, epiplectus and fresvillensis Zones age. As
pointed out above, Lewyites ambindense (Collignon) is

109

known only from the lower Upper Maastrichtian.

A specimen of Desmophyllites diphylloides (Forbes, 1846)
was found in bed 21 at Jebel Rawdah, section 2, well up in the
Simsima Formation. However, the species is not age diagnos-
tic, and its known range is Santonian to Upper Maastrichtian,
fresvillensis Zone, and possibly higher. However, from Jebel
Rawdah section 1, from scree almost certainly derived from
the top of bed 4, we found a specimen Brahmaites (Anabrah-
maites) vishnu (Forbes, 1846). The few well-dated specimens
of this species are Upper Maastrichtian, fresvillensis Zone,
and possibly younger in age.

In the basal transgressive calcarenite shell bed beneath the
sands and conglomerates of the Qahlah Formation at Jebel
Agabah a Glyptoxoceras sp. was recovered. However, the
genus is not age diagnostic and ranges from Santonian to
upper Upper Maastrichtian. At Jebel Buhays, section 2, a
Libycoceras sp. was found in the basal conglomerate of the
Qahlah Formation. Libycoceras species first appear in the
Upper Campanian and may range into the Lower Maastrich-
tian (Zaborski, 1982).

The basal beds of the Simsima Formation at Jebel Buhays,
section 1, yielded a specimen of Pachydiscus (P.) dossantoi
(Maury, 1930) which suggests a ‘mid-Maastrichtian’ age,
presumably not very different to the Loftusia-rich beds at
Jebel Huwayyah.

In conclusion then, the meagre ammonite data that we
have suggests that the basal transgressive sands and conglom-
erates are late Campanian to ‘mid-Maastrichtian’ in age,
whereas the main carbonate deposits of the Simsima Forma-
tion are of early Upper Maastrichtian, fresvillensis Zone, age.
At Jebel Huwayyah the Loftusia-rich horizons may be as old
as late Campanian or as young as early Upper Maastrichtian,
fresvillensis Zone. We do not know whether or not sedimen-
tation continued to the end of the Maastrichtian.

Inoceramid biostratigraphy. Inoceramid biostratigraphy has
been based on what appear to be evolving lineages, often
expressed as the ranges of subspecies. These are usually
defined as the timespan of particular ‘morphs’ within an
evolving species, although short-term biological events, such
as the widespread ‘flood’ of a particular species or subspecies,
are also important. Both methods can provide useful bios-
tratigraphical data.

In recent years much progress has been made on integrat-
ing biostratigraphic data derived from Upper Cretaceous
inoceramids and other fossil groups with data from sequence
stratigraphy. Of all the Upper Cretaceous stages, however,
the inoceramid biostratigraphic scheme for the Maastrichtian
is the least well-established.

Species of the genus Trochoceramus hold the most promise
for establishing a sound Maastrichtian biostratigraphy, since
they apparently represent a relatively simple evolutionary
lineage. Trochoceramus species have a somewhat rounded
shell outline, with their umbones set back from the anterior,
and have evenly spaced, rounded, comarginal ribs and dis-
tinctive radial ribs. The earliest species is narrow, with a
regularly curving shell, but later species have a more convex
shell, which is narrow in the early stages, but then undergoes
a significant change in coiling direction, markedly increasing
the relative volume of the mantle cavity. A similar change is
also observed in the earlier genus Cremnoceramus.

Here we recognize three successive inoceramid faunas of
Trochoceramus which can be used to zone the Maastrichtian.
The lower division is characterized by the narrow, regularly

110

curved species Trochoceramus cf. monticulae (Fugger &
Kastner) (?= Trochoceramus radiosus (Quaas)). These flat
forms first occur in very late Campanian strata (e.g. in the
Nacatoch Sand of Navarro County, Texas, U.S.A. (Stephen-
son 1941, pl. 13, fig. 3), and from Ammoniten Berg in the
western Egyptian desert, apparently co-occuring with Libyco-
ceras ismaeli (Quaas, 1902). However, the species becomes
much more widespread in the lower part of the Lower
Maastrichtian (Dhondt, 1983).

The middle division is characterized by a more convex
species of Trochoceramus, T. ianjonaensis (Sornay), its con-
vexity arising from a sharp change in shell curvature following
an initial flattish stage. Such forms are known from Madagas-
car, where they have been dated as ‘lower’ Maastrichtian
(Besairie 1972), although the ammonites listed suggest levels
well into the Upper Maastrichtian as defined here. They are
also known both from the Calabar district of Nigeria, where
they are ‘mid’-Maastrichtian, and the St. Lucia Formation of
Zululand, Republic of South Africa, where they almost
certainly extend into the Upper Maastrichtian. Finally, they
are also reported from Libya, but apparently not associated
with ammonites.

The upper division is characterized by Trochoceramus
morgani (Sornay). It is of Upper Maastrichtian age and, so
far, has only been recognized from the Calcaire a Baculites of
the Cotentin Peninsula, France. 7. morgani clearly differs
from T. ianjonaesis by the smaller average size of the initial
flat part of the shell (this average is 67% of the average size in
T. ianjonaensis). Put another way, T. ianjonaensis includes
forms with large and small initial shells, whereas 7. morgani
includes only specimens with small initial shells. Because this
reduction in size of the initial flat portion of the shell has not
been recognized in Trochoceramus from the Republic of
South Africa, it is uncertain whether this size reduction
represents a genuine world-wide change in the lineage or
simply a geographical variation. A tentative biostratigraphic
scheme for the Maastrichtian based on inoceramid bivalves
and compared with the cephalopod biochronology is pre-
sented in Fig. 12.

Although inoceramid bivalves are proving useful for the
division of the Maastrichtian they occur only rarely in the
Upper Cretaceous deposits of the U.A.E. and Oman. We
have discovered them at only three horizons, and so far only
the material from Jebel Rawdah is at all well-preserved.
Furthermore, none of this material includes species of Tro-
choceramus.

The lowest horizon includes only a fragment indeterminate
at generic level, which was recovered from the conglomerate
series below the Loftusia-beds at Jebel Huwayyah.

At the second horizon, in the basal transgressive shell bed
of the Simsima Formation at Jebel Bu Milh, we found a single
large fragmentary specimen of ‘Platyceramus’ sp. This genus
occurs in some abundance in the Maastrichtian succession of
the St Lucia Formation in the Republic of South Africa,
where it is common in the Lower Maastrichtian, but also
seems to be dominant in the uppermost inoceramid horizon
of that formation. A rather poorly preserved fragment of
Endocostea ct. bebahoaensis comes from the Loftusia-beds at
Jebel Huwayyah. It is not sufficiently well preserved to be
dated better than Campanian or Maastrichtian.

The highest of the three levels, some way up the Simsima
Formation at Jebel Rawdah, has yielded the most important
inoceramid fauna. At Jebel Rawdah, section 1, just below the
base of the more massive, well-cemented limestones (base of

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

bed 5) Endocostea (Cataceramus) semaili sp. nov. occurs.
This is most comparable with specimens from Nagoryany,
Ukraine, and indicates a ‘mid-Maastrichtian’ horizon. Drs
Skelton and Nolan also collected this species, together with
Endocostea (Endocostea) cf. coxi, Endocostea sp. and
‘Endocostea’ bebahoaensis, from the south-western part of
the jebel at the same locality and possibly the same horizon as
the ammonite Pachydiscus (P.) neubergicus neubergicus.
Endocostea coxi occurs with Trochoceramus ianjonaensis in
Nigeria and Zululand, while ‘Endocostea bebahoaensis
occurs with the same Trochoceramus in Madagascar, but is
known to be long-ranging (it occurs in the Upper Maastrich-
tian at Cotentin, France). The inoceramids from Jebel Raw-
dah imply an early Upper Maastrichtian age, thus giving
support to the age suggested by the ammonites from the same
horizon.

Echinoid biostratigraphy. Although useful biostratigraphi-
cally for division of late Cretaceous—Palaeocene strata in the
Boreal realm, echinoid distribution in Tethyan carbonate
sequences is at present too poorly known to provide reliable
dating of the succession. Furthermore, the environmental
constraints of most species makes them highly restricted in
their occurrence, and thus of limited value. However, our
work has identified two biostratigraphically useful species
lineages, Hemipneustes spp. and Hattopsis spp.

There are three species of Hemipneustes which differ in the
relative elevation of the test, sharpness of the anterior sulcus
and position of the apical disc. At Jebel Rawdah the lowest
beds contain the flattest species, H. persicus Cotteau and
Gauthier, and this species has a relatively broad and shallow
anterior sulcus. Above this level, and possibly co-occurring
for a short interval, comes H. arabicus, a species that is
equally flat in profile but with a narrower and more sharply
defined anterior sulcus. Higher beds contain an elevated
species of Hemipneustes, H. compressus, and the relative
elevation of the test increases up the succession, so that in the
upper half of the Simsima Formation, Hemipneustes compres-
sus begins to develop a strong apical elevation and narrower
frontal groove, resembling the boreal species H. striatoradi-
atus.

The oldest species, Hemipneustes persicus, occurs (as H.
sardanyolae Vidal) in the late Campanian of Spain. H.
arabicus is endemic to this area, but the highest species, H.
compressus, is known from the Maastrichtian of the Mari
Hills, West Pakistan, where, unfortunately, the precise dating
of the beds remains uncertain. At Jebel Rawdah, the replace-
ment of H. persicus by H. arabicus seems to take place within
the Upper Maastrichtian, according to the evidence of the
ammonites and inoceramids.

The second biostratigraphically useful lineage comprises
the two species of Hattopsis. At Jebel Faiyah, two species
succeed one another, with H. paucituberculatus predating H.
sphericus. This is useful for local correlation, since it is H.
paucituberculatus that occurs at the base of the Simsima
Formation at Jebel Buhays and H. sphericus that occurs near
the base of the Simsima Formation at Jebel Rawdah. This
then implies that the base of the Simsima Formation is not
strictly contemporaneous between jebels.

Finally, it is noteworthy that the basal bed in the Loftusia-
rich unit at Jebel Huwayyah yields a distinct suite of echinoids
seen at no other jebel, including the holectypoid Coptodiscus.
Coptodiscus is known from the late Campanian of Arabia

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

111

Rawdah 2

. roo
Faiyah Poh
[a0
cr rj See
Inferred sea-level change = oo
deep <> shallow ‘=o ‘aoe Rawdah 3
Humeayga Cot Coo
J] = ecos
fof =] 1S
saa se
roo
=o
aww

as Buhays ==
= ro
= ‘ot

Siliciclastic sands, grits and conglomerates

HE Poorly sorted bioclastic packstones

Poorly sorted orbitoid-rich packstones and wackestones

Well-sorted bioclastic packstones

Red-weathering silt-rich wackestone

Fig. 13 Tentative correlation between jebels with inferred sea-level curves.

| (Kier 1972) and from a presumed similar ‘Senonian’ horizon
/ in southern Iran.

Global Sea-level curves. In broad terms the stratigraphic

| Successions in the five jebels described here follow the same
pattern, but in detail, precise correlation remains difficult,
due to rapid shifts in facies and bed thickness across even
small areas (Figs 6, 13). Sedimentation upon the Semail
_Nappe and Hawasina Group follows a typical transgressive
pattern, often with massive boulder beds of serpentinite at
the base overlain by arenitic and then calc-arenitic sediments,
tall deposited in shallow or nearshore marine environments.
The lower clastic facies of conglomerates, sands and gravels
jare commonly included in the Qahlah Formation, whereas
tiie sediments above, which are essentially bioclastic cal-
Carenites, form the Simsima Formation. There is no reason to

suppose, however, that the change from boulder beds and
essentially quartz clastics upwards into calcarenites took
place at the same time in all the sections we have examined.

The general deepening and shallowing cycles that can be
recognized within individual sections allow a means of corre-
lation. In all sections the initial coarse clastics of the Qahlah
Formation are abruptly terminated, presumably marking the
submersion of the ophiolitic complex within the region. This
allowed carbonate production to dominate. For most of the
Simsima Formation deposition appears to have more-or-less
kept pace with subsidence so that sea-level remained around
wave-base.

Furthermore, at Jebel Buhays, Jebel Faiyah and Jebel
Rawdah there are conspicuous reddened beds and partings
that probably contain land-derived iron oxides, which are a
common constituent of lateritic soils formed on exposed

ophiolite. These red partings and thin beds presumably
formed when lateritic soils were eroded and periodically
flushed out to sea in run-off after heavy rains. Some may
mark minor fluctuations in sea-level. They are potentially
traceable over a wide area and may be important for local
correlation. The problem is that there are not the same
number of iron-enriched bands currently recognized in the
different sections, and it becomes problematic as to which of
several alternative bands should be correlated. A tentative
correlation is presented in Fig. 13. However, mineralogical
and geochemical analyses of these bands is needed to estab-
lish whether any have distinctive signatures.

The Cretaceous part of the section at Jebel Rawdah is
topped by a limestone conglomerate apparently made up of
eroded fragments of the Simsima Formation and including
large chunks of rudists. This is in turn overlain by Lower
Tertiary limestones. The lack of rounding and poor sorting of
the clasts in the limestone conglomerate are probably indica-
tive of subaerial erosion and deposition but we have no way
of more accurately dating this period of emergence. It lies
somewhere between late Maastrichtian and early Tertiary.

Flexer & Reyment (1989) identified two late Cretaceous
transgressive events affecting the Arabo-Nubian shield: one
in the late Campanian-early Maastrichtian, and the other in
the late Maastrichtian. Although local tectonic events, associ-
ated with the ophiolite emplacement, may have had a pro-
found effect on the local sea-level signature, it is tempting to
associate the initial submergence of the ophiolitic islands and
the sands and conglomerates of the Qahlah Formation with
the late Campanian-early Maastrichtian transgression, and
the later flooding of these beds and initiation of carbonate
platform deposition with the second of these major sea-level
transgressions in the late Maastrichtian. The end of the
Cretaceous saw a major drop in sea-level.

Summary. Combining the evidence from ammonites, inoc-
eramid bivalves, echinoids and global sea-level curves we
conclude that the basal siliciclastic beds of the Qahlah Forma-
tion are of latest Campanian age. At Jebel Huwayyah, the
Loftusia-rich levels of the Qahlah Formation probably
encompass Lower to early Upper Maastrichtian. Finally, the
Simsima Formation appears to be early Upper Maastrichtian,
fresvillensis Zone or later.

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APPENDIX

Macrofossils collected in the United Arab Emirates/Oman border region. Echinoids identified by A.B. Smith, nautiloids, bivalves and
gastropods by N.J. Morris, P.W. Skelton and R.J. Cleevely, ammonites by W.J. Kennedy, corals by J. Darrell and B.R. Rosen, bryozoans by
P.D. Taylor and brachiopods by E.F. Owen. Locality details are given in the main text: bed numbers refer to those shown in figures 5-11.
Numbers in square brackets after each name refer to the number of specimens collected.

Jebel Buhays, Section 1

Top of Bed 4. Echinoids: Arnaudaster cylindriformis sp. nov. [1];
Circopeltis? emiratus sp. nov. [1]. Ammonite: Pachydiscus dossan-
toi (Maury) [1].

Bed 10. Echinoids: Hemipneustes sp. [1]; Coenholectypus cf. baluchis-
tanensis (Noetling) [1].

Bed 11. Bivalves: Dictyoptychus morgani (Douvillé) [2]; Gastropods:
Acteonella crassa (Dujardin) [6].

Bed 12, base. Echinoids: Goniopygus arabicus sp. nov. [1]; Hattopsis
paucituberculatus sp. nov. [1 fragment]. Coral: Hydnophoraraea
sp. [1]. Bivalves: Scabrotrigonia sp. [3].

Bed 15. Echinoid: Codiopsis lehmannae sp. nov. [1]. Bivalve: Hippu-
rites cf. cornucopiae Defrance [6].

Lowest part of Simsima Formation (beds 4-10) — mostly collected

loose. Echinoids: Rhabdocidarid, gen. et sp. indet. [1]; Prionoci-
daris morgani (Gauthier) [2]; cidarid spines [2]; Heterodiadema
buhaysensis sp. nov. [2]; Orthopsis miliaris (d’ Archiac) [10]; Sale-
nia nutrix Peron & Gauthier [5]; Goniopygus arabicus sp. nov.
[22]; Glyphopneustes hattaensis Ali [41]; Hattopsis paucitubercula-
tus sp. nov. [11]; Noetlingaster paucituberculatus (Noetling) [3];
Phymosoma hexoaporum Lambert [13]; Actinophyma spectabile
Cotteau & Gauthier [4]; Plistophyma asiaticum Gauthier [3];
Circopeltis? emiratus sp. nov. [1]; Coenholectypus inflatus (Cotteau
& Gauthier) [8]; Coenholectypus cf. baluchistanensis (Noetling)
[3]; ‘Globator’ bleicheri (Thomas & Gauthier) [96]; Conulus dou-
villei (Cotteau & Gauthier) [19]; Vologesia rawdahensis Ali [3];
Pygurostoma morgani Cotteau & Gauthier [26]; Petalobrissus cf.
setifensis (Cotteau) [1]; Nucleopygus magnus sp. nov. [27]; Arnau-
daster cylindriformis sp. nov. [5]; Hemipneustes sp. indet. (frag-
ments) [3]; Hemiaster hattaensis Ali [4]. Bivalves: Arca sp. [1,
bivalved]; Cucullaea sp. A [3, 2 bivalved]; ‘Modiolus’ cf. capitatus
Zittel [3, bivalved]; Myzilus sp. nov? [1, bivalved]; Modiolus aff.
typicus Forbes [3, bivalved]; Neithea regularis (Schlotheim) [33];
Spondylus sp. C [1]; Spondylus sp. D [1]; Ctenoides aff. scaberrima
(Stolizcka) [14]; ?Osculopha sp. [1]; ?Gyropleura sp. [1]; ?Pycn-
odonte uncinella (Leymerie) [8]; Amphidonte pyrenaicum (Leyme-
tie) [2]; Agerostrea ungulata (Schlotheim) [4]; Eligmidae [6,

bivalved]; ?Tellinella sp. [1]; Glabrobournonia arabica Morris &
Skelton sp. nov. [12]; Tancrediidae cf. Tatella sp., sp. nov. [1];
Clavagella cf. semisulcata Forbes [2, bivalved]; Pholadomya sp. B
[1, bivalved]. Gastropods: Bathrotomaria cf. verdachellensis
(Forbes) [2]; Calliomphallus sp. [1]; Angaria sp. [1]; ‘Tectus’ ex.
gr. rozeti (d’Archiac) [7]; Umboniinae nov. gen. [9]; ?Rhabdocon-
cha sp. [1]; Turritellidae [2]; Pyrazus sp. [1]; Campanile sp. [8];
Tylostoma incerta (Forbes) [9]; Naticidae [1]; Strombidae, gen.
nov., cf. crassicostatus (Noetling) [1]; Strombidae, gen. nov. [1];
Strombacea [3]; ?Ovula expansa dArchiac & Haime [4];
Cypraeidae [1]; ?Tonnacea [1]; ‘Volutilithes’ dubia Noetling [7];
?Volutoderma sp. [4]; Caricella sp. [1]; Volutidae A [1]; Volutidae
B [2]; Volutidae C [2]; Volutidae D [3]; Neogastropoda indet. [1];
Acteonella crassa (Dujardin) [4]; Acteonella caucasica Zekeli cf.
styriaca Kollman [3]; Acteonella sp. cf. A. borneensis Nuttall &
Leong [19]; Acteonella caucasica Zekeli subsp. nov. [7]; Acteonella
crassa (Dujardin) [9]; Neocylindrites cf. minutus Sohl [3]; Neocylin-
drites sp. [1]; opisthobranch [2]. Corals: Polytremacis sp. [2]; Aspidas-
traea sp. [1]; Cunnolites sp. [4]; Diploctenium sp. [3]; Hydnophoraraea
sp. [2]; Paraplacocoenia orbignyana (Reuss) [1]; cerioid colony [1];
phaceloid colony [1]; solitary form [1]. Stromatoporoid: massive
stromatoporoid (bored by bivalves) [1]. Sponge: chaetetid [1]. Bryo-
zoan: ‘Onychocella’ sp. [1]. Decapod crustaceans: Carcineretes sp. [3
carapaces, 5 limb segments]; crab carapace, species A [1]; crab
carapace, species B [2]; claw, indet. [1].

Jebel Buhays Section la

Small hill 150 m to south of Jebel Buhays 1. Collected from scree, in
rock fall from lowest 4 m of Simsima Formation (beds are steeply
dipping and all loose material must be derived from the basal beds
here).

Echinoids: Heterodiadema buhaysensis sp. nov. [1]; Orthopsis
miliaris (d’Archiac) [7]; Salenia nutrix Peron & Gauthier [2];
Goniopygus arabicus sp. nov. [7]; Glyphopneustes hattaensis Ali [9];
Phymosoma hexoaporum Lambert [3]; Actinophyma spectabile Cot-
teau & Gauthier [1]; Plistophyma asiaticum Gauthier [3]; Coenholec-
typus inflatus (Cotteau & Gauthier) [3]; ‘Globator’ bleicheri (Thomas

114

& Gauthier) [occurs]; Conulus douvillei (Cotteau & Gauthier)
[occurs]; Vologesia rawdahensis Ali [2]; Petalobrissus linguiformis
(Peron & Gauthier) [1]; Nucleopygus magnus sp. nov. [9]; Hemip-
neustes sp. [1]; Hemiaster hattaensis Ali [3]. Bivalves: Cucullaea sp. A
[4]; Barbatia sp. [1]; Lyriochlamys ternatus (Minster) [6]; Neithea
regularis (Schlotheim) [6]; Agerostrea ungulata (Schlotheim) [4];
?Pycnodonte uncinella (Leymerie) [1, bivalved]; Scabrotrigonia sp.
[8]; Dictyoptychus morgani Douvillé [1, lid]; Glabrobournonia ara-
bica Morris & Skelton sp. nov. [2]; Clavagella cf. semisulcata Forbes
[2]. Gastropods: Tylostoma incerta (Forbes) [2]; ?Strombidae, gen.
nov. [1]; Pseudomelania [Trajanella] sp. cf. conica (Stolizcka) [2],
Acteonella crassa (Dujardin) [13]; Acteonella cf. laevis laevis (Sow-
erby) [6]; Acteonella cf. borneensis Nuttall and Leong [4]; Neocylin-
drites sp. cf minutus Stolizcka [1]. Cephalopods: Cimomia aff.
sowerbyana (d’Orbigny) [1]; Deltoidonautilus salisfilius sp. nov. [1].
Corals: Polytremacis sp. [1]; Favia sp. cf. ‘Diplocoenia’ klogsdorfen-
sis Trauth [1]; Barysmilia irregularis (Reuss) [1]; Cunnolites sp. [1];
Diploctenium sp. [1]; cerioid colony [1]. Sponge [1].

Jebel Buhays 2

Collections made from the scree-slope. Here the bedding is very
steep and material is derived from the lowest few metres of section
only.

Echinoids: Salenia nutrix Peron & Gauthier [1]; Goniopygus
arabicus sp. nov. [4]; Coenholectypus inflatus (Cotteau & Gauthier)
[1]; ‘Globator’ bleicheri (Thomas & Gauthier) [3]; Conulus douvillei
(Cotteau & Gauthier) [3]; Pygurostoma morgani Cotteau & Gauthier
[2]. Bivalves: Dictyoptychus morgani (Douvillé) [1]; Glabro-
bournonia arabica Morris & Skelton, sp. nov. [16]; Ammonite:
Libycoceras sp. [1].

Jebel Buhays 3

Basal limestone (lowest Im). Echinoids: Goniopygus arabicus sp. nov.
[4]; Nucleopygus magnus sp. nov. [1].

Above first major red-weathering siltstone level. Echinoids: Circopel-
tis emiratus sp. nov. [2].

Loose. Echinoids: Salenia nutrix Peron & Gauthier [2]; Hattopsis
paucituberculatus sp. nov. [2]; Phymosoma hexoaporum Lambert
[1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]. Bivalves: Hippu-
rites cornucopiae Defrance [1]; ?Biradiolites aff. baylei Toucas [2].

Jebel Thanais

Lowest 4 m of Simsima Formation. Echinoids: Heterodiadema buhay-
sensis sp. nov. [1]; Orthopsis miliaris (d’Archiac) [1]; Salenia nutrix
Peron & Gauthier [2]; Goniopygus arabicus sp. nov. [2]; Glyphop-
neustes hattaensis Ali [6]; Hattopsis sp. [1]; Phymosoma
hexoaporum Lambert [3]; Plistophyma asiaticum Gauthier [2];
Coenholectypus cf. baluchistanensis (Noetling) [2]; ‘Globator’ ble-
icheri (Thomas & Gauthier) [5]; Conulus douvillei (Cotteau &
Gauthier) [8]; Vologesia rawdahensis Ali [1]; Pygurostoma mor-
gani Cotteau & Gauthier [1]; Nucleopygus magnus sp. nov. [4];
Hemiaster hattaensis Ali [1]. Bivalves: ?Barbatia sp. A, cf. B.
morgani (Douvillé) [2, bivalved]; ‘Modiolus’ aff. typicus Forbes [1,
bivalved]; Neithea regularis (Schlotheim) [2]; Pycnodonte vesicu-
laris (Lamarck) [1, bivalved]. Gastropods: Acteonella sp. cf. A.
borneensis Nuttall & Leong [1]; Acteonella caucasica Zekeli subsp.
cf. styriaca Kollman [1]; Acteonella caucasica caucasica Zekeli [1].
Corals: Cunnolites sp. [1]; Diploctenium sp. [1]; Moltkia isis
Steenstrup [1]; Paraplacocoenia orbignyana (Reuss) [1]; coarse
meandroid colony [1]. Bryozoan: ‘Onychocella’ sp. [1]. Decapod
crustacean: limb segments [3].

Upper beds (equivalent to bed 15 of Jebel Buhays 1). Bivalves:
Hippurites cornucopiae Defrance [1].

Loose in scree. Bivalves: Cucullaea sp. A [2 ,1 bivalved]; ‘Modiolus’

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

aff. typicus Forbes [1, bivalved]; Chlamys dujardeni (Roemer) [1];
Neithea regularis (Schlotheim) [3]; Spondylus sp. E [1]; Ctenoides
aff. scaberrima (Stolizcka) [2]; ?Amphidonte cf. pyrenaicum
(Leymerie) [1]; ?Osculopha sp. [1]; Lucinidae gen. indet., sp. B [1,
bivalved]; Dictyoptychus morgani (Douvillé) [1]; Glabrobournonia
arabica Morris & Skelton, sp. nov. [1]; Colveraia variabilis Kling-
hardt [1]; Vaccinites vesiculosus (Woodward) [4]; Lapeirousia sp.
[2]; Pholadomya sp. B [1, bivalved]. Gastropods: Tylostoma incerta
(Forbes) [1]; Acteonella crassa (Dujardin) [5].

Jebel Aqabah

Excellent outcrop of the basal part of the marine sequence resting
directly on serpentinized ultramafics.

Bed 1. Echinoids: Hattopsis paucituberculatus sp. nov. [1]; ‘Globator’
bleicheri (Thomas & Gauthier) [2]; Faujasia eccentripora Lees [6
juveniles]. Ammonite: Glyptotoxoceras sp. [1].

Loose. Bivalves: ?Pycnodonte uncinella (Leymerie) [2]; Dictyopty-
chus morgani (Douvillé) [1]; ?Biradiolites aff. baylei Toucas [3].
Gastropods: Acteonella crassa (Dujardin) [2]. Cephalopod: Cimo-
mia aff. sowerbyana (d’Orbigny) [1]. Coral: Cunnolites sp. [1].

Jebel Huwayyah, Section 1

Bed 1. Bivalves: ?Acutostrea sp. [ca. 10].

Bed 3. Bivalves: ?Acutosrea sp. [abundant].

Bed 7. Stromatoporoid [1].

Bed 9. Echinoids: Faujasia eccentripora Lees [6]; Hemiaster sp. cf. H.
hattaensis Ali [1]. Bivalves: Spondylus sp. A [1]; Vaccinites vesicu-
losus (Woodward) [5]; ?Granocardium sp. [1]. Gastropods: ?Tec-
tus sp. [1]; Campanile sp. [1]; Amauropsina aff. bulbiformis (J. de
C. Sowerby) [1]; ‘Amputllina aff. splendida’ (Deshayes) [3]. Corals:
Cunnolites sp. [4]; cerioid colony (with Lithophaga borings) [1].
Ammonite: Pachydiscus sp. [1].

Beds 10-11. Echinoids: Faujasia eccentripora Lees [33]; Hemiaster sp.
cf. H. hattaensis Ali [1]. Bivalves: Endocostea cf. bebahoaensis
(Sornay) [1]; Spondylus sp. A [2]; Plicatula hirsuta Coquand [4];
?Acutostrea sp. [6]; Glabrobournonia arabica Morris & Skelton,
sp. nov. [1]; Pholadomya? sp. B [1]. Gastropods: cf. Tylostoma
incerta (Forbes) [1]. Corals: Cunnolites sp. [2]; Paraplacocoenia
orbignyana (Reuss) [1]; large trochoid solitary [1]; large flabellate
solitary [1]; cerioid colonies (5 genera) [7]. Ammonites: Pachydis-
cus dossantoi (Maury) [1]; Lewyites ambindense (Collignon) [1];
Nostoceras (Nostoceras) major Kennedy & Cobban [1].

Beds 9-11 undifferentiated. Bivalves: Torreites sanchezi (Douvillé) [2]
Durania form A [1].

Bed 13. Bivalves: Durania spp. [fragments].

Beds 14/15. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier)
[1]; ‘Globator’ bleicheri (Thomas & Gauthier) [54]; Pygurostoma
morgani Cotteau & Gauthier [3]; Hemipneustes sp. [1]. Bivalves:

Cucullaea sp. A [1]; Lyriochlamys ternatus (Minster) [7]; Neithea |

regularis (Schlotheim) [8]; Spondylus sp. A. [2]; Dictyoptychus

morgani (Douvillé) [1]. Gastropods: Execochirsus sp. [1]; Campa- |

nile sp. [1]; Cerithiidae [1]; Tylostoma incerta (Forbes) [6]; Strom-
bacea [1]; ?Aporrhaidae [1]; Neogastropods [2]. Ammonite:
Pachydiscus dossantoi (Maury) [1]. Coral: Cunnolites sp. [1].

Bed 16. Bivalves: Pycnodonte vesicularis (Lamarck) [2]; Agerostrea
ungulata (Schlotheim) [12, bivalved].

Bed 17. Bivalves: Lyriochlamys ternatus (Minster) [1]; Neithea
regularis (Schlotheim) [2]; Scabrotrigonia sp. [4]; Durania sp. 2].
Gastropods: Bathrotomaria sp. [1].

Bed 18, towards top of section. Echinoids: Proraster geayi Cottreau
{7].

Loose. Echinoid: Mecaster sp.? [1]. Bivalve: ?‘Inoceramus’ sp. [1].

———————
- ————————

|
|

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

Jebel Huwayyah, Section 2

Bed 1. Echinoids: G/yphopneustes hattaensis Ali [1]; Coptodiscus

magniproctus sp. nov. [2].

Beds 3-5. Echinoids: cidarid spine [1]; Orthopsis sp. [1]; Salenia

microprocta sp. nov. [1]; ?Hattopsis sp. [1]; Faujasia eccentripora
Lees [1]; Hemiaster sp. cf. H. hattaensis Ali [3]; Mecaster sp. [4];
Proraster geayi Cottreau [2]. Bivalves: Dictyoptychus morgani
(Douvillé) [1]; Corals: Cunnolites sp. [27]; Diploctenium sp. [4];
Placosmilia sp. [22]. Bryozoans: cf. Euritina lata Canu [1]; ‘Wilber-
topora’ sp. [1]. Decapod crustacean: pincer [1]. Brachiopods:
terebratulid gen. et sp. nov. [3]. Ammonite: Nostoceras sp. [1].

Bed 7. Bivalves: Vaccinites vesiculosus (Woodward); Corals: Actina-

cis sp. [3]; Cladocora ?humilis (Michelin) [7]; ?Aspidastraea sp. [1];
Astraraea sp. [3]; Calamophylliopsis simonyi (Reuss) [1]; Parapla-
cocoenia orbignyana (Reuss) [1]; cerioid colonies (3 genera) [4];
thamnasteroid colonies [4]; coarse meandroid colonies [5];
elongate-oval flat solitary corals [3]; flabellate solitary corals [2];
turbate solitary corals [10]; cylindrical solitary corals [3]. Ammo-
nite: Neancycloceras sp. [1].

Loose in scree, but derived from Loftusia beds, beds 2—7 unless

otherwise stated. Echinoids: Prionocidaris? emiratus sp. nov. [1];
Coenholectypus inflatus (Cotteau & Gauthier) [3] (probably
derived from Simsima Formation above?). Bivalves: Endocostea
cf. bebahoaensis (Sornay) [1]; Lyriochlamys ternatus (Minster) [8];
Neithea regularis (Schlotheim) [19]; Neithea (Neithella) notabilis
(Munster) [1]; Spondylus sp. A [49, 19 bivalved]; Spondylus sp. B
[12, 4 bivalved]; Plicatula hirsuta Coquand [23]; ?Osculopha sp. [4,
1 bivalved]; Amphidonte pyrenaicum (Leymerie) [10, 2 bivalved];
Agerostrea ungulata (Schlotheim) [1, bivalved]; ?Eligmidae indet.
[1]; Ctenoides sp. [2]; Chama noetlingi [7]; Biradiolites aff. baylei
Toucas [1]; Glabrobournonia arabica Morris & Skelton, sp. nov.
[10]; Semailia smithi Morris & Skelton, sp. nov. [1]; Vaccinites
vesiculosus (Woodward) [25]; Pholadomya sp. A, cf. indica Noet-
ling [2]; Pholadomya sp. B [1, bivalved]. Gastropods: ?Angaria sp.
[5]; Turbinidae sp. [5]; Turritellidae sp. [10]; ?Pyrazus sp. [3];
Cerithtidae sp. [1]; ?Perrisoptera sp. [7]; ?Naticidae sp. [7]; ?Fasci-
olariidae sp. [1]; Volutidae sp. A [1]; Volutidae sp. B [1];
?Mitridae sp. [1]; neogastropods [2]; Acteonella sp. [1].

| Jebel Rawdah, Section 1

Bed 2. Echinoid: Petalobrissus cf. setifensis (Cotteau) [1]. Bivalves:

?*Modiolus’ cf. capitatus Zittel [1, bivalved]; Cucullaea sp. A [11];
Lyriochlamys ternatus (Minster) [2]; Neithea regularis (Schlo-
theim) [7]; Pycnodonte vesicularis (Lamarck) [1]; ?Pycnodonte
uncinella (Leymerie) [7]; Agerostrea ungulata (Schlotheim) [1];
Indet. ribbed oyster [1]; Scabrotrigonia sp. [62, 7 bivalved];
Durania spp. [2]; Glabrobournonia arabica Morris & Skelton [4];
Tancrediidae, cf. Tatella, sp. nov. [2, bivalved]. Gastropods:
Acteonella sp. cf. borneensis Nuttall & Leong [4]. Corals: Cunno-
lites sp. [7]; Diploctenium sp. [1].

Bed 3 (lower part) and top of Bed 2. Echinoids: Orthopsis miliaris

(@Archiac) [2]; Salenia nutrix Peron & Gauthier [2]; Hattopsis
sphericus Ali [19]; Noetlingaster paucituberculatus (Noetling) [5,
fragments]; Coenholectypus cf. baluchistanensis (Noetling) [1];
‘Globator’ bleicheri (Thomas & Gauthier) [49]; Faujasia eccentri-
pora Lees [5]; Zuffardia morgani (Cotteau & Gauthier) [4];
Petalobrissus cf. setifensis (Cotteau) [17]; Phymechinus? sp. [frag-
ment]. Bivalves: ?Barbatia sp. A, cf. B. morgani Douvillé [1,
bivalved]; Cucullaea sp. A [4, 3 bivalved]; ?‘Mytilus’ nitens Forbes
[1, bivalved]; ‘Modiolus’ cf. capitatus Zittel [5, 4 bivalved]; Pinna

_ sp. [3]; Endocostea (Selenoceramus) semaili sp. nov. [1]; Lyrio-

chlamys ternatus (Minster) [3]; Neithea regularis (Schlotheim)
[23]; Limidae [1]; Plicatula hirsuta Coquand [1]; Pycnodonte
vesicularis (Lamarck) [6]; Pycnodonte vesicularis (Lamarck) var.
hippopodium Nillson [1]; ?Pycnodonte uncinella (Leymerie) [10, 4

115

bivalved]; ?Amphidonte cf. pyrenaicum (Leymerie) [2]; Scabrot-
rigonia sp. [6, 2 bivalved]; Lucinidae sp. [1]; ?Plagyoptychus sp.
[1]; Dictyoptychus morgani (Douvillé) [10]; Glabrobournonia ara-
bica Morris & Skelton, sp. nov. [22]; Durania spp. [6 fragments];
?Trapeziidae [1]; ?Mesocallista sp. [5, bivalved]; Tellininae [1];
?Tancrediidae, cf. Tatella, sp. nov. [2]; Clavagella cf. semisulcata
Forbes [6]; ?Brechites cf. aspergilloides (Forbes) [1]; Pholadomya
sp. B [1]. Gastropods: Calliomphalus sp. [4]; ?Angaria sp. [2];
?Tectus sp. [3]; ?Pseudoliotina sp. [1]; ?Umboniinae, gen. nov.
[64]; euomphalid gen. nov. [6]; ?Rhabdoconcha sp. [1]; ?Turritel-
lidae [2]; Vermetidae or Siliquariidae [1]; Cerithiidae [7];
Cypraeidae, sp. [4]; ?Cypraeidae. [3]; ?Volutacea [1]; Ampullina
aff. ‘splendida’ (Deshayes) [4]; Tylostoma incerta (Forbes) [2];
Mesogastropod [2]; ?Anchura sp. [3]; 7Aporrhaidae [1]; Hipponi-
cidae [3]; ?Fasciolariidae [1]; ?Volutidae [16]; ?Buccinacea [7];
Lyria sp. [8]; ?Lyria sp. [2]; Acteonella sp. cf. borneensis Nuttall &
Leong [30]; Acteonella laevis laevis (J. de C. Sowerby) [1];
Acteonella sp. [4]; Neocylindrites sp. [2]; oliviform opisthobranch
[1]; ?Scaphander sp. [1]; bullaeform opisthobranchs [2]; elongate
opisthobranch [1]. Cephalopods: Cimomia cf. sowerbyana
(d’Orbigny) [1]; Brahmaites (Anabrahmaites) vishnu (Forbes) [1].
Corals: Cunnolites sp. [42]; Diploctenium sp. [9]; Aspidastraea sp.
[2]. Decapod crustacean: pagurid claw [1].

Bed 3 (upper part). Echinoids: Hattopsis sphericus Ali [3]; phymo-

somatid fragment [1]; Conulus douvillei (Cotteau & Gauthier) [1];
Zuffardia morgani (Cotteau & Gauthier) [1]; Hemipneustes sp. [3,
fragments]. Bivalves: ?Chlamys sp. [1]; ?Ctenoides sp. [1]; Cardi-
idae [3]; ?Fragum sp. [2]. Gastropods: ?Umboniinae, gen. nov. [6];
?Rhabdoconcha sp. [2]; Campanile sp. [1]; ?Cerithiidae indet. [3];
?‘Ampullina aff. splendida’ (Deshayes) [1]; ?Tylostoma incerta
(Forbes) [4]; ?Naticidae sp. [2]; Cypreidae spp. [3]; Lyria sp. [2];
elongate, fusiform genus [1].

Bed 4. Echinoids: Glyphopneustes hattaensis Ali [1]; Hattopsis spheri-

cus Ali [15]; Hemipneustes arabicus Ali [3]. Bivalves: Neithea sp.
[1]; Durania form B [1]; ?Pycnodonte uncinella (Leymerie) [7]; cf.
Fragum sp. [3]. Gastropods: ?Angaria sp. [1]; ?Trochacea,
euomphalid gen. nov. [3]; ?Umboniinae, gen. nov. [4]; Turritel-
lidae [1]; ?Cerithiidae, sp. [2]; Cypraeidae, sp. [1]; Caricella sp.
[2]. Cephalopod: Cimonia aff. sowerbyana (d’Orbigny) [1]. Deca-
pod crustacean: small spinose pincers [2].

Bed 4 (top). Echinoids: Salenia nutrix Peron & Gauthier [2]; Hattopsis

sphericus Ali [29]; Noetlingaster emiratescus Ali [1]; Noetlingaster
sp. [1, fragment]; Phymechinus perplexus sp. nov. [2]; ‘Globator’
bleicheri (Thomas & Gauthier) [15]; Faujasia eccentripora Lees [2];
Hemipneustes persicus Cotteau & Gauthier [1, fragment]. Bivalves:
?‘Mytilus’ nitens Forbes [1, bivalved]; ‘Modiolus’ aff. typicus
Forbes [2, bivalved]; Endocostea (Selenoceramus) semaili sp. nov.
[2]; Neithea sp. [6]; ?Pycnodonte uncinella (Leymerie) [16]; Sca-
brotrigonia sp. [5, 1 bivalved]; Gyropleura sp. [4]; ?Tellinella sp.
[1, bivalved]; ‘Quenstedtiidae’, gen. nov. [1]; Clavagellidae, gen.
nov. [3]. Gastropods: ?Umboniinae, gen. nov. [27]; Turritellidae
[1]; ?Rhabdoconcha sp. [2]; Cerithiidae [1]; Strombidae, gen. nov.
[3]; Buccinacea sp. [1]; Lyria sp. [2]; ?Volutomorpha sp. [2];
Volutidae, cf. Melo [1]; ?Cancellariidae [1]; 7Neogastropods indet.
[5]. Cephalopod: Cimonia cf. sowerbyana (d’Orbigny) [1]. Corals:
Cunnolites sp. [4].

Collected in scree or from slipped blocks. Echinoids: Orthopsis

miliaris (d’Archiac) [4]; Salenia nutrix Peron & Gauthier [1];
Noetlingaster sp. [1, fragment]; ‘Globator’ bleicheri (Thomas &
Gauthier) [2]; Faujasia eccentripora Lees [6]; Petalobrissus sp. [2];
Hemipneustes compressus Noetling [1]; Hemipneustes sp. [2, frag-
ments]. Bivalves: Cucullaea sp. A [2, 1 bivalved]; Neithea regularis
(Schlotheim) [1]; Pycnodonte vesicularis (Lamarck) [2]; ?Pycn-
odonte uncinella (Leymerie) [2]; Agerostrea ungulata (Schlotheim)
[3]; Scabrotrigonia sp. [14]; Glabrobournonia arabica Morris &
Skelton, sp. nov. [1]; Radiolitidae [1]; Tancrediidae, cf. Tatella, sp.
nov. [1, bivalved]. Gastropods: ‘Tectus’ sp. [1]; Turbinidae [1];

116

?2Umboniinae, gen. nov. [1]; ?Turritellidae indet. [1]; ‘Ampullina
aff. splendida’ (Deshayes) [2]; Tylostoma incerta (Forbes) [2];
2Naticidae sp. [1]; ?Cimolithium sp. [1]; ?Aporrhaidae sp. [2];
Lyria sp. [3]; Acteonella sp. cf. borneensis Nuttall & Leong [8];
opisthobranch [1]. Corals: Cunnolites sp. [66]; Diploctenium sp.
[1]. Ammonites: Brahmaites (Anabrahmaites) vishnu (Forbes) [1].

Bed 6. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [8];
‘Globator’ bleicheri (Thomas & Gauthier) [1]; Hemipneustes perst-
cus Cotteau & Gauthier [2]; Hemiaster hattaensis Ali [3].

Rebel Rawdah, Section 2

Bed 4: 60-120 cm above basal conglomerate. Echinoids: Goniopygus
arabicus sp. nov. [1]; Plistophyma asiaticum Gauthier [1]; Echino-
tiara perebaskinei Lambert [6]; Faujasia eccentripora Lees [14];
Petalobrissus rawdahensis sp. nov. [3]. Bivalve: Durania [1]. Gas-
tropod: ?‘Ampullina aff. splendida’ (Deshayes) {2]. Decapod crus-
tacean: limb segments [4].

Bed 5. Echinoids: Faujasia eccentripora Lees [2].

Beds 6-8. Echinoids: Salenia nutrix Peron & Gauthier [10]; Goniopy-
gus arabicus sp. nov. [6]; Codiopsis lehmanni sp. nov. [1]; Phy-
mechinus perplexus sp. nov. [6]; ?Phymechinus sp. [1]; Echinotiara
perebaskinei Lambert [58]; Coenholectypus inflatus (Cotteau &
Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1];
Faujasia eccentripora Lees [92]; Petalobrissus rawdahensis sp. nov.
[4]; Petalobrissus cf. setifensis (Cotteau) [9]; Hemipneustes arabicus
Ali [1]. Bivalve: ?Ctenoides sp. [1]; Durania form B [4]. Coral:
Cunnolites sp. [1]. Decapod crustacean: limb and claw segments
[4].

Bed 10. Echinoids: Goniopygus arabicus sp. nov. [1]; Glyphopneustes
hattaensis Ali [2]; Faujasia eccentripora Lees [1]; Petalobrissus sp.
[2]. Bivalves: Durania cf. apula (Parona) [1]. Gastropod: Stromba-
cea [1].

Bed 11: Cucullaea bed immediately overlying orange-weathering
band. Echinoids: Noetlingaster paucituberculatus (Noetling) [1,
fragment].

Bed 11 (middle, ca. 1 m above orange-weathering band). Echinoids:
Goniopygus arabicus sp. nov. [1]; Faujasia eccentripora Lees [6];
Petalobrissus sp. [1]; Nucleopygus magnus sp. nov. [4]; Stigmatopy-
gus pulchellus sp. nov. [2]; Hemipneustes arabicus Ali [4]. Gastro-
pods: ?Umboniinae, gen. nov. [1]; Campanile sp. [1]; Pyrazus sp.
[1]; Strombidae, gen. nov. [1]; Volutidae sp. [3].

Bed 11: 2.5 m above base. Echinoids: Echinotiara perebaskinet
Lambert [18]; Coenholectypus cf. baluchistanensis (Noetling) [1];
Faujasia eccentripora Lees [49]; Petalobrissus sp. [40]. Gastropod:
‘Ampullina aff. splendida’ (Deshayes) [1]. Corals: Cunnolites sp.
[3]; massive meandroid colony [1].

Bed 11 (unspecified). Echinoids: Salenia nutrix Peron & Gauthier [2];
Glyphopneustes emiratensis Ali [5]; Hattopsis sphericus Ali [2];
Noetlingaster paucituberculatus (Noetling) [4]; Phymechinus? per-
plexus sp. nov. [1]; Echinotiara perebaskinei Lambert [3]; Coen-
holectypus cf. baluchistanensis (Noetling) [8]; ‘“Globator’ bleicheri
(Thomas & Gauthier) [10]; Faujasia eccentripora Lees [46]; Zuffar-
dia morgani (Cotteau & Gauthier) [41]; Petalobrissus rawdahensis
sp. nov. [247]; Petalobrissus cf. setifensis (Cotteau) [124]. Bivalves:
‘Modiolus’ aff. typicus Forbes [1, bivalved]; Cucullaea sp. A [8,
bivalved]; Pholadomya sp. C. cf. P. connectans Forbes [22,
bivalved]. Coral: Cunnolites sp. [1].

Bed 13. Echinoids: Salenia nutrix Peron & Gauthier [2]; Glyphop-
neustes hattaensis Ali [1]; ‘Globator’ bleicheri (Thomas &
Gauthier) [1]; Petalobrissus rawdahensis sp. nov. [230]; Petalobris-
sus cf. setifensis (Cotteau) [4]; Nucleopygus magnus sp. nov. [3].
Bivalves: ‘Modiolus’ aff. typicus Forbes [1, bivalved]; Dictyopty-
chus morgani (Douvillé) [common]; ?Biradiolites aff. baylei Tou-
cas [1]; ?Fragum sp. [1]; ?Clavagellidae indet. [1]. Gastropods:
?Umboniinae, gen. nov. [6]; Turritellidae, sp. [1]; ?Rhabdoconcha
sp. [1]; ?}Cimolithium sp. [2]; ‘Ampullina aff. splendida’ (Deshayes)

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

Bed 14. Echinoids: Noetlingaster paucituberculatus (Noetling) [5 plus

Bed 15. Echinoids: Conulus douvillei (Cotteau & Gauthier) [2];

Beds 16-19. Bivalve: ‘Modiolus’ aff. typicus Forbes [1, bivalved].

Bed 16. Echinoids: ‘Globator’ bleicherei (Thomas & Gauthier) [3];

Bed 18. Echinoid: Pygurostoma morgani Cotteau & Gauthier [1].

Bed 19. Echinoids: Noetlingaster paucituberculatus (Noetling) [4];

Bed 20. Echinoids: ‘Globator’ bleicheri (Thomas & Gauthier) [2];

Bed 21 (base). Echinoids: Noetlingaster paucituberculatus (Noetling

[4]; Cypraeidae, sp. [1]; cf. ?Columbellidae [2]; ?Neogastropod
indet. [1]. Corals: Cunnolites sp. [15]; Diploctenium sp. [2];
Aspidastraea sp. {1}.

fragments]; Phymechinus ? sp. [1]; ‘Globator’ bleicheri (Thomas &
Gauthier) [7]; Conulus douvillei (Cotteau & Gauthier) [1]; Fauja-
sia eccentripora Lees [7]; Pygurostoma morgani Cotteau &
Gauthier [3]; Arnaudaster cylindriformis sp. nov. [4]; Petalobrissus
rawdahensis sp. nov. [17]; Petalobrissus cf. setifensis (Cotteau)
[15]; Petalobrissus linguiformis (Peron & Gauthier) [4]; ?Stigmato-
pygus pulchellus sp. nov. [6]; Hemipneustes arabicus Ali [1]; H.
compressus Noetling [2]. Gastropods: Campanile sp. [2]; ?Campa-
nile sp. [1]; ‘Ampullina aff. splendida’ (Deshayes) [2]; ? Pseudocas-
sis sp. [1]; Strombidae, gen. nov. [2]; Fasciculariidae, sp. [1];
?Volutidae indet. [1]. Corals: Aspidastraea sp. [1].

Petalobrissus rawdahensis sp. nov. [25]; Hemipneustes compressus
Noetling [2]. Bivalves: ?Granocardium sp. [3]; rudist [1]. Gastro-
pods: Pyrazus sp. [2]; Cerithiidae [1]; ?Naticidae, sp. [1]; Strom-
bidae gen. nov. [3]. Corals: Cunnolites sp. [21]; Aspidastraea sp.
[9]; massive meandroid colony [1].

Corals: Cunnolites sp. [2]; Aspidastraea sp. (2].

Faujasia eccentripora Lees [3]; Petalobrissus rawdahensis sp. nov.
[5]. Bivalves: Neithea regularis (Schlotheim) [1] Durania spp.
[several]; Veneridae [2]. Gastropods: ?Rhabdoconcha sp. [I];
?Cerithiidae [1]; ‘Ampullina’ aff. splendida (Deshayes) [3]; Strom-
bidae [1]; Cypraeidae [1].

Bivalves: ?Granocardium sp. [1]. Gastropods: ?Umboniinae gen.
nov. [1]; ?Cimolithium sp. [2]; Strombidae, gen. nov. [1].

Coenholectypus inflatus (Cotteau & Gauthier) [1]; ‘Globator’
bleicheri (Thomas & Gauthier) [8]; Vologesia rawdahensis Ali [1];
Zuffardia morgani (Cotteau & Gauthier) [5]; Faujasia eccentripora
Lees [19]; Arnaudaster cylindriformis sp. nov. [1]; Petalobrissus
rawdahensis sp. nov. [73]; Petalobrissus cf. setifensis (Cotteau)
[23]; Petalobrissus linguiformis (Peron & Gauthier) [5]; ?Stigmato-
pygus pulchellus sp. nov. [1]; Pygurostoma morgani Cotteau &
Gauthier [2]; Hemipneustes compressus Noetling [1]; Hemipneustes
persicus (Cotteau & Gauthier) [1]. Bivalves: ?}Granocardium sp.
[2]; Isognomon sp. [1]; ?Lucinidae indet. [1]; Dictyoptychus mor-

gani (Douvillé) [1]; Veneridae, gen. indet. [1]. Gastropods:)

?Umboniinae, gen. nov. [9]; ?Rhabdoconcha sp. [1]; ?Cerithiidae
[5]; Campanile sp. [4]; ‘Ampullina aff. splendida’ (Deshayes) [3);
Tylostoma incerta (Forbes) [2]; ?Naticidae [4]; Strombidae, gen.
nov. [5]; Cypraeidae, sp. [1]; Fasciolariidae or Buccinidae [1];

Canalicella sp. [1]; Volutidae [3]; Acteonella sp. [1]. Corals:|

Neocaeniopsis reussi (Edwards & Haime) [1]; Astrogyra edwardsi

(Reuss) [1]; cerioid colonies [2]; Cunnolites sp. [17]; Diploctentum| |

sp. [1]; Aspidastraea sp. [8]; massive meandrine colonies [5].

Faujasia eccentripora Lees [2]; Petalobrissus linguiformis (Peron &

Gauthier) [1]; Petalobrissus sp. [1]; Hemipneustes compressus

Noetling [1]; Hemipneustes persicus (Cotteau & Gauthier) [1]

Gastropods: ?Angaria sp. [1]; ?Umboniinae gen. nov. [2]:
‘Ampullina aff. splendida’ (Deshayes) [3]; Strombidae, gen. nov.
[5].

[1]; ‘Globator’ bleicheri (Thomas & Gauthier) [1]; Conulus douvil
lei (Cotteau & Gauthier) [2]; Faujasia eccentripora Lees [22]
Zuffardia morgani (Cotteau & Gauthier) [5]; Pygurostoma mor
gani Cotteau & Gauthier [7]; Arnaudaster cylindriformis sp. NOV
[1]; Petalobrissus rawdahensis sp. nov. [45]; Petalobrissus cf
setifensis (Cotteau) [44]; Petalobrissus linguiformis (Peron

Gauthier) [4]; ?Stigmatopygus pulchellus sp. nov. [7]; Hemip

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

neustes compressus Noetling [16]; Hemipneustes persicus (Cotteau
& Gauthier) [22]. Bivalves: Barbatia cf. morgani (Douvillé) [3,
bivalved]; Pycnodonte vesicularis (Lamarck) [1]; heterodont indet.
[1]. Gastropods: Campanile sp. [1]; ‘Ampullina aff. splendida’
(Deshayes) [1]; Naticidae [2]; Tylostoma incerta (Forbes) [1];
Strombidae, gen. nov. [1]; ?Fasciolariidae [1]; bullaeform opistho-
branch [1]. Ammonite: Desmophyllites diphylloides (Forbes) [1].

Bed 21 (middle). Gastropods: ‘Ampullina aff. splendida’ (Deshayes)
[4]; Strombidae, gen. nov. [3]; Strombidae, sp. [1]; ?Hercorhyn-
chus sp. [1]; ?Buccinacea, sp. [1]; neogastropod, gen. A [2];
neogastropod, gen. B [1].

Bed 22. Echinoids: ‘Globator’ bleicheri (Thomas & Gauthier) [1];
Faujasia eccentripora Lees [3]; Petalobrissus rawdahensis sp. nov.
[13]; Hemipneustes compressus Noetling [1]. Gastropods:
‘Ampullina aff. splendida’ (Deshayes) [2]; ?Buccinacea [1].

Bed 23 (base). Echinoids: Petalobrissus sp. [1]. Gastropods: Pyrazus
sp. [1]; Naticidae [1].

Beds 23-25. Echinoids: Petalobrissus sp. [5]; Faujasia eccentripora
Lees [1]. Bivalves: ? Biradiolites aff. baylei Toucas [2].

Bed 25. Echinoids: Noetlingaster paucituberculatus (Noetling) [4];
Faujasia eccentripora Lees [3]; Zuffardia morgani (Cotteau &
Gauthier) [1]; Petalobrissus sp. [11]. Bivalve: ?Lucinidae indet. [1].

Bed 26. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1];
Arnaudaster cylindriformis sp. nov. [1]; Pygurostoma morgani
Cotteau & Gauthier [2]; Petalobrissus rawdahensis sp. nov. [5];
Petalobrissus cf. setifensis (Cotteau) [3]; Hemipneustes compressus
Noetling [2]; Hemipneustes sp. fragment [1]. Bivalve: Neithea sp.
{1].

' Bed 27. Echinoids: Hemipneustes compressus Noetling [2].

/ Bed 28. Bivalves: Indet. small rudists.

Loose: a little below beds 21/22 and probably derived from them.
Echinoids: Hemipneustes arabicus Ali [1].

Loose: scree from level of bed 11 (derived from beds 13-15 mostly).
| Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1];
Conulus douvillei (Cotteau & Gauthier) [1]; Faujasia eccentripora
Lees [14]; Zuffardia morgani (Cotteau & Gauthier) [3]; Petalobris-
sus linguiformis (Peron & Gauthier) [6]; Petalobrissus sp. [2].

| Loose, scree. Echinoids: Glyphopneustes hattaensis Ali [2]; Echinoti-
ara perebaskinei Lambert [2]; Coenholectypus inflatus (Cotteau &
| Gauthier) [1]; ‘Globator’ bleicheri (Thomas & Gauthier) [4];
| Faujasia eccentripora Lees [32]; Zuffardia morgani (Cotteau &
| Gauthier) [11]; Arnaudaster cylindriformis sp. nov. [1]; Pyguros-
| toma morgani Cotteau & Gauthier [1]; Petalobrissus sp. [62];
| Nucleopygus magnus sp. nov. [3]; Hemipneustes persicus (Cotteau
| & Gauthier) [4]; Hemiaster hattaensis Ali [1]. Bivalves: ‘Modiolus’
| aff. typicus Forbes [1, bivalved]; Spondylus sp. C [1]; Amphidonte

pyrenaicum (Leymerie) [1]; Dictyoptychus morgani (Douvillé) [1];
Radiolites sp. [1]; ?Biradiolites aff. baylei Toucas [2]. Gastropods:
?Umboniinae, gen. nov. [3]; Pyrazus sp. [1]; Campanile sp. [1];
‘Ampullina aff. splendida’ (Deshayes) [1]; ?Perrisoptera sp. [1];
Caricella sp. [2]; Acteonella sp. [1]; Acteonella cf. caucasica Zekeli
| [1]. Corals: Cunnolites sp. [12]; Aspidastraea sp. [1].
oose, lower part of section (from beds 1-11). Echinoids: Salenia
nutrix Peron & Gauthier [3]; Goniopygus arabicus sp. nov. [4];
Glyphopneustes hattaensis Ali [4]; Hattopsis sphericus Ali [1];
Noetlingaster paucituberculatus Noetling [1]; Phymechinus? per-
plexus sp. nov. [4]; Echinotiara perebaskinei Lambert [15]; Coen-
holectypus baluchistanensis Noetling [5]; ‘Globator’ bleicheri
(Thomas & Gauthier) [9]; Faujasia eccentripora Lees [80]; Zuffar-
dia morgani (Cotteau & Gauthier) [29]; Petalobrissus spp. [136].
Asteroid: marginal [1]. Brachiopod: terebatulid, gen. nov. [1].
ose, mid-section. Echinoids: Salenia sp. [1]; Zuffardia morgani
(Cotteau & Gauthier) [6]; ?Stigmatopygus pulchellus sp. nov. [1].
Corals: Cunnolites sp. [2]; Aspidastraea sp. [1].

117
Jebel Rawdah, Section 3a

Bed 2. Echinoids: Goniopygus arabicus sp. nov. [2]. Bivalve: Durania
sp. [1]. Gastropod: Acteonella sp. [1].

Bed 4. Echinoid: Echinotiara perebaskinei Lambert [1]. Bivalves:
Pycnodonte vesicularis (Lamarck) [1]; Amphidonte pyrenaicum
(Leymerie) [2, bivalved].

Bed 5. Bivalves: ‘Modiolus’ aff. typicus Forbes [1, bivalved]; ?*Myzi-
lus’ nitens Forbes [1, bivalved]; Barbatia sp. B [3]; Neithea regularis
(Schlotheim) [3]; Amphidonte pyrenaicum (Leymerie) [3]; Ageros-
trea ungulata (Schlotheim) [9]; Eligmidae [5, bivalved]; Crassitel-
lites sp. [5].

Bed 7 (near top). Echinoid: ‘Globator’ bleicheri (Thomas & Gauthier)
[1]. Bivalves: Pycnodonte vesicularis (Lamarck) [1]; Agerostrea
ungulata (Schlotheim) [1, bivalved]; ?Clavagella sp. [tubes]. Gas-
tropods: Naticidae, indet. [2].

Loose, derived from lower beds. Bivalves: Plicatula hirsuta Coquand
[1]; ?Pycnodonte uncinella (Leymerie) [2, bivalved]. Gastropod:
Turritellidae [1].

Jebel Rawdah, Section 3b

Bed 2 (lower part). Echinoids: Orthopsis miliaris (d’Archiac) [1];
Glyphopneustes hattaensis Ali [1]; Circopeltis emiratus sp. nov. [1];
Faujasia eccentripora Lees [3]. Coral: Cunnolites sp. [1]. Decapod
crustacean: Callianassa (limb segment) [1].

Bed 2 (upper part). Echinoids: Salenia nutrix Peron & Gauthier [1];
Glyphopneustes hattaensis Ali [1]; ‘Globator’ bleicheri (Thomas &
Gauthier) [11]; Petalobrissus rawdahensis sp. nov [19]; Petalobris-
sus cf. setifensis (Cotteau) [4]; Nucleopygus magnus sp. nov. [3].
Coral: ?Diploctenium sp. [1].

Bed 3. Echinoid: Echinotiara perebaskinei Lambert [1]. Bivalve:
Pycnodonte vesicularis (Lamarck) [1].

Bed 4. Corals: large meandrine colonies [2]; placoid colony [1].

Bed 5. Echinoids: Heterodiadema buhaysensis sp. nov. [1]; Glyphop-
neustes hattaensis Ali [1]; Coenholectypus inflatus (Cotteau &
Gauthier) [3]; ‘Globator’ bleicheri (Thomas & Gauthier) [1];
Hemipneustes sp. [1]; Hemiaster hattaensis Ali [1]. Bivalve: Pycn-
odonte vesicularis (Lamarck) [1].

Bed 5/6. Echinoids: ‘Globator’ bleicheri (Thomas & Gauthier) [8];
Hemipneustes persicus Cotteau & Gauthier [1]; Mecaster victoris
Lambert [47].

Bed 6. Echinoid: Circopeltis emiratus sp. nov. [1]. Bivalve: Eligmidae
[1]. Gastropod: Tylostoma incerta (Forbes) [1].

Bed 7. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1];
Hemipneustes persicus Cotteau & Gauthier [1]. Bivalves: Barbatia
sp. B [1, bivalved)]; ‘Modiolus’ aff. typicus Forbes [2, bivalved].

Bed 8. Echinoids: Orthopsis miliaris (d’Archiac) [1]; Actinophyma
spectabile Cotteau & Gauthier [1]; Coenholectypus sp. [1]; Vologe-
sia rawdahensis Ali [1]; Faujasia eccentripora Lees [1]; Mecaster sp.
[6]. Coral: Cunnolites sp. [1]. Decapod crustacean: pincer [1].

Bed 9. Echinoids: Orthopsis miliaris (d’ Archiac) [7]; Noetlingaster sp.
[1, fragment]; Actinophyma spectabile Cotteau & Gauthier [1];
‘Globator’ bleicheri (Thomas & Gauthier) [1]; Faujasia eccentri-
pora Lees [3]; Hemipneustes compressus Noetling [2]; Hemiaster
sp. cf. H. hattaensis Ali [1]; Mecaster victoris Lambert [1]; Pro-
raster geayi Cottreau [2]. Bivalves: Cucullaea sp. A [4]; Scabrot-
rigonia sp. [8, 1 bivalved]; ?Tancrediidae, sp. nov. [1]. Gastropods:
Acteonella sp. cf. borneensis Nuttall & Leong [1]; Acteonella sp.
[2]. Corals: Cunnolites sp. [8]; Diploctenium sp. [1]; thamnasteroid
colony [1].

Bed 11. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1];
‘Globator’ bleicheri (Thomas & Gauthier) [1]; Arnaudaster cylin-
driformis sp. nov. [1]. Bivalves: Barbatia sp. B [2]; ‘Modiolus’ aff.
typicus Forbes [1, bivalved]; ? Pycnodonte uncinella (Leymerie) [3,
bivalved]; Amphidonte pyrenaicum (Leymerie) [2, 1 bivalved];

118

Agerostrea ungulata (Schlotheim) [21]; Scabrotrigonia sp. [7, 1
bivalved]. Gastropod: Caricella sp. [1].

Loose. Echinoids: Prionocidaris morgani (Gauthier) [1]; Orthopsis
miliaris (d’Archiac) [3]; Coenholectypus inflatus (Cotteau &
Gauthier) [6]; ‘Globator’ bleicheri (Thomas & Gauthier) [3];
Conulus douvillei (Cotteau & Gauthier) [4]; Hemipneustes com-
pressus Noetling [1].

Loose near top of section. Echinoids: Arnaudaster cylindriformis sp.
nov. [1]; ?Linthia sudanensis (Bather) [1].

Jebel Rawdah, Section 4

Bed 1. Bivalves: Dictyoptychus morgani (Douvillé) [1]; Praeradiolites
cf. subtoucasi Toucas [4]; Pseudosabinia aff. klinghardti Boehm
[3]; Pseudosabinia sp. [1].

Bed 1/2. Echinoids: Phymosoma hexoaporum Lambert [1]; ‘Globator’
bleicheri (Thomas & Gauthier) [6]. Bivalves: Scabrotrigonia sp. [2];
Glabrobournonia arabica Morris & Skelton, sp. nov. [21].

Bed 2 (mostly near top). Echinoids: Echinotiara perebaskinei Lambert
[1]; Coenholectypus sp. indet. [1]; ‘Globator’ bleicheri (Thomas &
Gauthier) [3]; Faujasia eccentripora Lees [1]; Zuffardia morgani
(Cotteau & Gauthier) [2]; Petalobrissus cf. setifensis (Peron &
Gauthier) [1]; Hemiaster hattaensis Ali [1]. Bivalves: Neithea
regularis (Schlotheim) [1]; Pycnodonte vesicularis (Lamarck) [1];
?Pycnodonte uncinella (Leymerie) [3, 1 bivalved]; Amphidonte
pyrenaicum (Leymerie) [1]; Scabrotrigonia sp. [1]. Gastropod:
Acteonella sp. cf. borneensis Nuttall & Leong [1].

Bed 4. Echinoids: Glyphopneustes hattaensis Ali [1]; Circopeltis
emiratus sp. nov. [1]; ‘Globator’ bleicheri (Thomas & Gauthier)
[4]; Faujasia eccentripora Lees [2]; Petalobrissus sp. [1]. Corals:
Cunnolites sp. |2|; Aspidastraea sp. [1].

Bed 5. Echinoid: ‘Globator’ bleicheri (Thomas & Gauthier) [1].

Bed 8. Echinoids: Orthopsis miliaris (d’Archiac) [1]; Echinotiara
perebaskinei Lambert [1]; ‘Globator’ bleicheri (Thomas &
Gauthier) [3]; Conulus douvillei (Cotteau & Gauthier) [1]; Petalo-
brissus cf. setifensis (Cotteau) [3].

Bed 9. Bivalves: Amphidonte pyrenaicum (Leymerie) [1]; Scabrot-
rigonia sp. [common].

Bed 10. Echinoids: Orthopsis miliaris (d’ Archiac) [6]; Salenia nutrix
Peron & Gauthier [1]; ‘Globator’ bleicheri (Thomas & Gauthier)
[10]. Bivalve: Dictyoptychus sp. [1]. Corals: cerioid colony [1];
placoid colony [1].

Bed 12. Echinoids: Coenholectypus sp. [1]; ‘Globator’ bleicheri (Tho-
mas & Gauthier) [5]; Nucleopygus magnus sp. nov. [1]. Sponge:
chaetetid [1].

Bed 13. Echinoids: Noetlingaster emiratescus Ali [2]; Coenholectypus
inflatus (Cotteau & Gauthier) [1]; ‘Globator’ bleicheri (Thomas &
Gauthier) [3]; Conulus douvillei (Cotteau & Gauthier) [1]; Hemi-
aster hattaensis Ali [1].

Bed 15. Echinoids: Orthopsis miliaris (d’Archiac) [1]; ‘Globator’
beleicheri (Thomas & Gauthier) [1]. Coral: Actinacis sp. [1].

Bed 18. Echinoid: Coenholectypus sp. [1].

Bed 19. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier) [1];
‘Globator’ bleicheri (Thomas & Gauthier) [1]; Pygurostoma mor-
gani Cotteau & Gauthier [1].

Bed 22/23. Echinoids: Coenholectypus inflatus (Cotteau & Gauthier)
[1]; Faujasia eccentripora Lees {1].

30 cm below top of measured section. Echinoids: Coenholectypus
inflatus (Cotteau & Gauthier) [1]. Bivalve: Neithea regularis
(Schlotheim) [1].

Loose, a little below the top of the section. Echinoids: Hemipneustes
arabicus Ali [1].

Loose at level of bed 3. Echinoid: Petalobrissus linguiformis (Peron &
Gauthier) [1].

£.oose. Echinoids: Orthopsis miliaris (d’ Archiac) [1]; Conulus douvil-
lei (Cotteau & Gauthier) [1]; Petalobrissus sp. [1]. Bivalves:
Amphidonte pyrenaicum (Leymerie) [2].

A.B. SMITH, N.J. MORRIS, W.J. KENNEDY AND A.S. GALE

Jebel Faiyah, Section la

Bed 6. Bivalves: Hippurites aff. lapeirousei Goldfuss [1 colony].

Bed 8. Bivalves: Agerostrea ungulata (Schlotheim) [2]; ?Pycnodonte
uncinella (Leymerie) [18]; ?Amphidonte cf. pyrenaicum (Leyme-
rie) [1].

Jebel Faiyah, Section 1b

Bed 2 (lower part). Echinoids: Hattopsis paucituberculatus sp. nov.
[6]. Corals: Polytremacis sp. [1]; Astraraea sp. [1]; Hydno-
phoraraea sp. [1]. Sponge: chaetetid [1].

Bed 2 (upper part). Echinoids: cidarid [1]; Orthopsis miliaris
(d’Archiac) [1]; Salenia nutrix Peron & Gauthier [2]; Glyphop-
neustes hattaensis Ali [1]; Hattopsis paucituberculatus sp. nov. [1];
‘Globator’ bleicheri (Thomas & Gauthier) [1]; Nucleopygus mag-
nus sp. nov. [1]; Hemiaster sp. cf. H. hattaensis Ali [1]. Bivalves:
Hippurites cf. lapeirousei Goldfuss [1 colony]; Hippurites aff.
cornucopiae Defrance [1]; Dictyoptychus morgani (Douvillé) [2];
Biradiolites aff. baylei Toucas [1]; Durania cf. gaensis (Dacqué)
[1]. Decapod crustacean: pincers [2].

Bed 4. Bivalves: Hippurites aff. lapeirousei Goldfuss [2]. Corals:
Polytremacis sp. [1]; Hydnophoraraea sp. [1]; Paraplacocoenia
orbignyana (Reuss) [1]; Moltkia isis Sreenstrup [1].

Bed 5. Echinoids: Glyphopneustes hattaensis Ali [2].

Bed 6. Echinoid: cidarid spine [1]. Coral: Moltkia isis Sreenstrup [1].

Bed 7. Echinoids: cidarid spine [1]; Mimiosalenia quinquetuberculata
sp. nov. [21]; Hattopsis sphericus Ali [1]; ‘Globator’ bleicheri
(Thomas & Gauthier) [36].

Bed 8. Coral: Cunnolites sp. [1].

Loose (derived from beds 2-5). Echinoids: Hattopsis paucituberculatus
sp. nov. [5]. Bivalves: Spondylus sp. C [2]. Nautiloid: Deltoidonau-
tilus salisfilius sp. nov. {1]. Corals: Polytremacis sp. [5]; Cunnolites
sp. [1]; Paraplacocoenia orbignyana (Reuss) [1]; Hydnophoraraea
sp. [5]; turbinate solitary [1]; cerioid colonies (6 genera) [14];
flabellate solitary [1]; phaceloid colony [1]. Branching algae [1].
Sponge: chaetetid [1]. Decapod crustacean: crab carapace [1].

Jebel Faiyah — southern nose of Jebel (bed
numbers as in section la)

Bed 8. Echinoids: Salenia sp. [1]; Glyphopneustes hattaensis Ali [8];|
Hattopsis sphericus Ali [8]; ‘Globator’ bleicheri (Thomas &
Gauthier) [2]; Pygurostoma morgani Cotteau & Gauthier [1].
Corals: Cunnolites sp. [2].

Loose. Echinoids: Hattopsis sphericus Ali [1]. Bivalves: ?Barbatia sp.
A. cf. B. morgani (Douvillé) [1, bivalved]; Neithea regularis
(Schlotheim) [7]; ?Pycnodonte uncinella (Leymerie) [18]; ?Pla-
gioptychus sp. [1]; Glabrobournonia arabica Morris & Skelton, sp.
nov. [1]; Hippurites 2cornucopiae Defrance [11]; Dictyoptychus
morgani (Douvillé) [3]. Gastropod: Tylostoma incerta (Forbes) [1].
Corals: Moltkia isis Streenstrup [1]; Cunnolites sp. [3]; Hydno-
phoraraea sp. [2]; Paraplacocoenia orbignyana (Reuss) [1]; cerioid
colonies (3 genera) [4]; phlacoid colony [1]; thamnasteroid colony

[1].

Jebel Faiyah, Section 2

Basal 1 m shell bed. Bivalves: Glabrobournonia arabica Morris &
Skelton, sp. nov. [3]. Nautiloids: Cimonia cf. sowerbyan
(d’Orbigny) [1]; Deltoidonautilus salisfilius sp. nov. [2].

Jebel Bu Milh, Section 1
Bed 1. Bivalves: Pseudosabinia aff. klinghardti (Boehm) [4]; Durant

cf. gaensis (Dacqué).
Bed 3. Bivalves: ‘Modiolus’ aff. typicus Forbes [1, bivalved]; Lyrio

LATE CRETACEOUS CARBONATE PLATFORM FAUNAS

chlamys ternatus (Minster) [2]; Neithea regularis (Schlotheim) [1];
?Plagioptychus sp. [4, lids]; Dictyoptychus morgani (Douvillé) [1,
juvenile]; Radiolites sp. [3]; Eodictyoptychus aff. arumaensis Skel-
ton & El-Asa’ad [2]. Gastropods: ?‘Angaria’/Liotia sp. [1, frag-
ment]; ?trochid [1]; Discotectus sp. [26]; Strigosella sp. cf. striolata
(Stolizcka) [2]; Euchelus ornatus Stolizcka [3]; ?Umbonium greyi
Lees [1]; ‘Turritella’ sp. [1]; ‘Turritella’ sp. 1 ?= Nairiella multi-
striata (Reuss) [4]; Vermetus sp. [2]; Campanile curtum Douvillé
[1]; Campanile persicum Douvillé [2]; Campanile aff. breve Lees
[1, fragment]; Campanile morgani Douvillé [1]; Campanile sp. [6,
internal moulds]; Campanile cf. ganesha Noetling [4]; Cimolithium
sp. nov. [2]; Pyrazus sp. [1]; Paryphostoma morgani Douvillé [1];
Pugnellus sp. [2]; Ampullina aff. splendida (Deshayes) [30+];
‘Euspira’ lirata J. de C. Sowerby [22]; ‘Natica’ pagoda Forbes [2];
Tylostoma incerta (Forbes) [5]; Confusiscala sp. [1]; ‘Cypraea’
kayeii Forbes [2]; Hipponyx sp. [1]; ‘Fulguraria’ multistriata Stolic-
zka [1]; fasciolariid [1]; ?Rapanidae gen. indet. [1]; ‘Murex’ sp. [1];
‘Trophon’ oldhamianum Stolizcka [1]; Volutoderma elongata
Stolizcka [1]; Volutolithes latisepta Stoliczka [2]; Caricella pyrifor-
mis Forbes [2]; Caricella sp. [2]; ‘Voluta’ sp. or spp. [3]; Trochac-
taeon sp. [1]; Acteonella cf. caucasica Zekeli [30+]; Acteonella
caucasica Zekeli subsp. grossouvrei Cossmann [10]; Acteonella
laevis laevis (J. de C. Sowerby) [4]; Acteonella elongata Kollmann
[4]. Corals: Aspidastraea sp. [1]; Hydnophoraraea sp. [2].

Jebel Bu Milh, Section 2

Bed 4. Gastropods: Acteonella cf. caucasica Zekeli [74]; Acteonella

elongata Kollmann [1]; Acteonella laevis (J. de C. Sowerby) [3];
Acteonella cf. laevis zekelii Kollmann [1]. Bivalves: Lapeirousia sp.

[2].

Beds 7/8. Echinoids: Petalobrissus cf. setifensis (Cotteau) [1].

eae

Bivalves: Barbatia sp. B [12, 4 bivalved]; Cucullaea sp. A [2,
bivalved]; Glycymeris sp. [1]; ‘Platyceramus’ sp. [1]; Lyriochlamys
ternatus (Munster) [1]; Neithea regularis (Schlotheim) [1]; Spondy-
lus sp. [4, bivalved]; Plicatula hirsuta Coquand [5]; Plagiostoma sp.
[2]; Crassatellites sp. [5]; Eodictyoptychus aff. arumaensis Skelton
& El-Asa’ad [1]; Biradiolites aff. baylei Toucas [1]. Gastropods:
Calliomphalus sp. [2]; ?Calliomphalus or Helicanthus sp. [1];
?Angaria’ sp. [2]; Cyclostrematid/Liotid gen. nov. [1]; cf. ‘Turbo’
punctatus Zekeli [1, fragment]; Discotectus sp. 1 [common]; Disco-
tectus sp. 2 [common]; Strigosella sp. cf. striolata (Stolizcka)
[common]; Nerita spp. [occurs]; ‘Lissocheilus’ persicus (Douvillé)
[4]; ‘Turritella’ sp. 1 ?= Nairiella multistriata (Reuss) [7]; ‘Turri-
tella’ sp. cf. morgani Douvillé [1]; ‘Turritella’ sp. 3 (2= Roemeriella
nerinea Akopyan (non Roémer)) [2]; Campanile ?robustum Dou-

119

villé [1]; Campanile curtum Douvillé [26]; Campanile ?persicum
Douvillé [2]; Campanile aff. breve (Lees) [5]; Campanile morgani
Douvillé [1]; Cimolithium sp. nov. [72]; Semivertagus cf. arcotense
Stolizcka [1]; cf. Pyrazella (Plicopyrazus) [1]; Pyrazus pyramidatus
Douvillé [1]; Exechocirsus sp. 1 [2]; Exechocirsus sp. 2 [1];
?Exechocirsus sp. [2]; ?Semivertagus sp. cf. arcotense (Stolizka)
[7]; Hantkenia louristana Douvillé [5]; Strombidae gen. nov.
giganteus Noetling [6]; Strombidae gen. nov. digitatus Noetling [2];
‘Helicaulax’ sp. [4]; Ampullina aff. splendida (Deshayes) [100];
‘Euspira lirata’ (J. de C. Sowerby) [1]; ?Tylostoma incerta (Forbes)
[12]; ‘Confusiscala’ sp. cf. turbinata Forbes [5]; ‘Calyptraea’ elevata
Forbes [1]; Thlacodes lamellosus Stolizcka [1]; Lathyrus sp. [1];
Lathyrus sp. cf. ‘Ornopsis’ digressa (Wade) [1]; ?Bellifusus sp. [1];
?Pseudoliva sp. [2]; ‘Voluta’ citharina Forbes [1]; Volutoderma sp.
[2]; Caricella pyriformis Forbes [20+]; ‘Voluta’ sp. [11]; ‘Voluta’
sp. [21]; Voluta cameleo Forbes [1]; Volutoderma sp. [1]; Lyria
crassicostata Dujardin [2]; ?Naronia eximia Stolizcka [1];
Acteonella caucasica Zekeli grossouvrei Cossmann [46]; Acteonella
caucasica caucasica Zekeli [31]; Acteonella caucasica Zekeli subsp.
nov. [1]; Acteonella laevis laevis (J. de C. Sowerby) [3];
?Acteonella laevis J. de C. Sowerby zekelii Kollman [1];
?Acteonella elongata Kollman [4]; Neocylindrites minutus
(Stolizcka) [2]. Ammonite: Nostoceras (Nostoceras) major
Kennedy & Cobban [1]. Nautiloids: Deltoidonautilus salisfilius sp.
nov. [9]. Corals: Cunnolites sp. [6]; cerioid colonies (2 genera) [3].
Sponges [7].
Bed 10. Bivalves: Vaccinites oppeli (Douvillé) [abundant, in situ].

Qarn Murrah

Rudist level near top of section. Bivalves: Osculigera cf. vautrinioides
Vogel [32]; Vaccinites vesiculosus (Woodward) [13];?Vaccinites
loftusi (Woodward) [3]; Glabrobournonia arabica Morris & Skel-
ton, sp. noy. [16]; Pseudosabinia aff. klinghardti (Boehm) [1];
Pseudopolyconites aff. parvus Milovanovic [1]; Colveraia variabilis
Klinghardt [1]. Corals: Cunnolites sp. [2]; cerioid colonies (2
genera) [3].

Qarn Mulayh

Lower part. Bivalves: Pironaea cf. polystylus Pirona [3]; Durania
form A [2]; Durania sp. [1].

7-7 m below the top of the section. Bivalyes: Torreites sanchezi
(Douvillé) milovanovici Grubic [3].

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Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):121—240

Issued 30 November 1995

Late Campanian-Maastrichtian echinoids
from the United Arab Emirates-Oman border

region

ANDREW B. SMITH

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD

CONTENTS
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Relative;abundance Of Species mrsrrn.sceeee eens as cect See nici’ -pysaits cesiems cclng se ste ete ole curse mer tee eeetneiors aoe ne neieeistem ceases waa 124
Echinoidipalacoccolopy, «ces, -ceceermc deceaet en sas geese ne ceeleathsnsiieadagh cas sreitmonsaateeoaepeeeeeaonteceseeoaueeesoet eh oeccee one 124
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alacoecologicalisymthesis; .. -aereemaceetane cess seettciae > a wae skool iolaccbistwaree wee aleealne ceria eemtietee tae maniac coca aslede seeee oaige 130
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IRELETENCES | cposcsta ist oa das teteee MOSSE Soe ee Ne «Sele ec wc eee Doiswienas ois» wa oe cetea Ot crate OM Pinee Sele ee eee teeta eink Stages ocean ge 239

Synopsis. Forty-five echinoid species, 14 of them new, are described from the Maastrichtian Simsima Formation

exposed along the western margins of the North Oman Mountains. The stratigraphic distribution of over 2,500
individual fossil echinoids has been recorded and used to assess echinoid abundance quantitatively and identify
recurrent assemblages of species. Although approximately equal numbers of regular and irregular echinoid species
are known, irregular echinoids greatly outnumber regulars in abundance. Furthermore, most species are rare, with
just five making up more than 67% of the total collection. Regular echinoids are divisible into four ecological
groups, ranging from hard-ground dwellers living within active wave-surge environments, to herbivores living in
protected subtidal soft-bottom environments. Five ecological groups of irregular echinoid are distinguished, largely
on the basis of their inferred feeding strategies. Seven echinoid assemblages are distinguished and their
palaeoecological setting interpreted on the basis of the autecology of included species, associated macrofauna and
lithofacies evidence. One new genus of Goniopygidae, Mimiosalenia, is described and the following new species are
erected: Prionocidaris? emiratus, Heterodiadema buhaysensis, Salenia microprocta, Goniopygus arabicus, Mimios-
alenia quinquetuberculata, Codiopsis lehmannae, Hattopsis paucituberculatus, Circopeltis? emiratus, Phymechinus?
perplexus, Coptodiscus magniproctus, Petalobrissus rawdahensis, Nucleopygus magnus, Stigmatopygus? pulchellus,

Arnaudaster cylindriformis.

INTRODUCTION

_ Late Cretaceous echinoid faunas, though well-documented in
_ both Europe and America, remain relatively little studied
| elsewhere in the world. Those of the Middle East are
particularly poorly known. Much of our knowledge of
Tethyan late Cretaceous echinoid faunas comes from the
| major monographic works of the last century such as those
dealing with Algeria (Cotteau et al. 1881), Iran (Cotteau &
Gauthier 1895; Gauthier 1902) and Baluchistan (Noetling
1897), all of which are now in great need of revision and
updating. Additional major late Cretaceous faunas were
| described during the early part of this century from Libya
_(Checchia-Rispoli 1930, 1931a,b, 1932a,b, 1933) and from
) Madagascar (Cottreau 1908; Lambert 1933).
_ Until recently the late Cretaceous faunas of the Arabian
'Peninsula remained virtually unknown. Duncan (1865)
‘described a few Cenomanian echinoids from south eastern
Oman, while Lees (1928) described two new late Cretaceous

©The Natural History Museum, 1995

echinoid species from north western Oman. Clegg (1933)
described a large number of echinoids from the Persian Gulf,
but only two of these were Cretaceous, and neither appears
to be late Cretaceous in age.

The first indication of the rich late Cretaceous echinoid
fauna of the Arabian Peninsula came with the publication of
Kier’s (1972) monograph on the Mesozoic and Tertiary
echinoids around Riyadh, Saudi Arabia. In this work he
described 11 species from the Campanian-Maastrichtian
Aruma Formation, six of which were new to science and the
remainder representing species already described by Cotteau
& Gauthier (1895) and Gauthier (1902) from southern Iran.
Subsequently, Ali (1989, 1992a,b) documented 20 echinoids,
including seven new species, from the Simsima and Qahlah
Formations of the western Oman Mountains. Smith (in
Skelton et al. 1990) gave some preliminary records of the
echinoid fauna from these beds and Roman et al. (1989) gave
a preliminary report on the Cretaceous echinoids of the
Dhofar region, Oman, including four Campanian/
Maastrichtian species.

A.B. SMITH

Cassiduloids

122

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Regular echinoid species

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Species (see Table 1 for key)
Emirates-Oman border region (based on ca. 2,500 individuals). Species 1—45 are listed in Table 1. Shading distinguishes the four major

taxonomic groups of echinoids. A, species arranged systematically. B, species arranged in order of specimen abundance. Pie diagrams

Fig. 1 Relative abundance of species collected from the Maastrichtian Qahlah and Simsima Formations along the United Arab
represent the number of individuals and number of species for each major taxonomic group.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

123

Table 1. Echinoid species collected from the Qalah and Simsima Formations exposed in the Jebels along the Oman-United Arab Emirates

border region.

Class Echinoidea Leske, 1778
Subclass Cidaroidea Claus, 1880
Order Cidaroida Claus, 1880
Family Rhabdocidaridae Lambert, 1900
1. Gen. et sp. indet.
Family Cidaridae Gray, 1825
Genus Prionocidaris Agassiz, 1863
Ds Prionocidaris morgani (Gauthier, 1902)
3. Prionocidaris? emiratus sp. nov.
Subclass Euechinoidea Bronn, 1860
?Cohort Diadematacea Duncan, 1889
?Order Diadematoida Duncan, 1889
Family Heterodiadematidae Smith & Wright, 1993
Genus Heterodiadema Cotteau, 1864
4. Heterodiadema buhaysensis sp. nov.
Cohort Echinacea Claus, 1876
Plesion (Order) Orthopsida Mortensen, 1942
Family Orthopsidae Duncan, 1889
Genus Orthopsis Cotteau, 1864
5. Orthopsis miliaris (d’ Archiac, 1835)
Order Calycina Gregory, 1900
Family Saleniidae Agassiz, 1838
Genus Salenia Gray, 1835
Salenia nutrix Peron & Gauthier, 1881
Salenia microprocta sp. nov.
Order Arbacioida Gregory, 1900
Family Goniopygidae Smith & Wright, 1993
Genus Goniopygus Agassiz, 1838
8. Goniopygus arabicus sp. nov.
Genus Mimiosalenia gen. nov.
9. Mimiosalenia quinquetuberculata sp. nov.
Family Glyphopneustidae Smith & Wright, 1993
Genus Glyphopneustes Pomel, 1883
10. Glyphopneustes hattaensis Ali, 1992

Family Arbactidae Gray, 1835
Genus Codiopsis Agassiz, 1840

11. Codiopsis lehmannae sp. nov.
Genus Hattopsis Ali, 1992
12. Hattopsis sphericus Ali, 1992
| 3. Hattopsis paucituberculatus sp. nov.
Genus Noetlingaster Vredenberg, 1911
(«14 Noetlingaster paucituberculatus (Noetling, 1897)
| iS. Noetlingaster emiratescus Ali, 1989
16. Noetlingaster? sp.

Order Phymosomatoida Mortensen, 1904
Family Phymosomatidae Pomel, 1883
Genus Phymosoma Haime, 1853

if. Phymosoma hexoaporum Lambert, 1927
| Genus Actinophyma Cotteau & Gauthier, 1895
| 18. Actinophyma spectabile Cotteau & Gauthier, 1895
Genus Plistophyma Peron & Gauthier, 1881
19: Plistophyma asiaticum Cotteau & Gauthier, 1895

Family Stomechinidae Pomel, 1883
Genus Circopeltis Pomel, 1883

f 20. Circopeltis? emiratus sp. nov.

; Genus Phymechinus Desor, 1856

| 21. Phymechinus? perplexus sp. nov.

Genus Echinotiara Pomel, 1883

22.

23.
24.

25.

26.

27.

28.

29.

30.

31.

32.
33.
34.

36.

35.

37.

45.

Echinotiara perebaskinei Lambert, 1929

Cohort Irregularia Latreille, 1825

Order Holectypoida Duncan, 1889
Family Holectypidae Lambert, 1899
Genus Coenholectypus Pomel, 1883
Coenholectypus inflatus (Cotteau & Gauthier, 1895)
Coenholectypus cf. baluchistanensis (Noetling, 1897)
Genus Coptodiscus Cotteau & Gauthier, 1895
Coptodiscus magniproctus sp. nov.
Family Conulidae Lambert, 1911
Genus ‘Globator’ Agassiz, 1840
‘Globator’ bleicheri (Gauthier, 1889)
Genus Conulus Leske, 1778
Conulus douvillei (Cotteau & Gauthier, 1895)
Order Cassiduloida Claus, 1880
Family Clypeolampadidae Kier, 1962
Genus Vologesia Cotteau & Gauthier, 1895
Vologesia rawdahensis Ali, 1989
Family Faujasiidae Lambert, 1905
Genus Faujasia d’Orbigny, 1856
Faujasia eccentripora Lees, 1928
Genus Zuffardia Checchia-Rispoli, 1917
Zuffardia morgani (Cotteau & Gauthier, 1895)
Unnamed family
Genus Pygurostoma Cotteau & Gauthier, 1895
Pygurostoma morgani Cotteau & Gauthier, 1895
Family Cassidulidae Agassiz & Desor, 1847
Genus Petalobrissus Lambert, 1916
Petalobrissus rawdahensis sp. nov.
Petalobrissus cf. setifensis (Cotteau, 1866)
Petalobrissus linguiformis (Peron & Gauthier, 1881)
Genus Stigmatopygus d’Orbigny, 1856
Stigmatopygus? pulchellus sp. nov.
Genus Nucleopygus Agassiz, 1840
Nucleopygus magnus sp. nov.
Family Echinolampadidae Gray, 1851
Genus Arnaudaster Lambert, 1918
Arnaudaster cylindriformis sp. nov.
Order Holasteroida Durham & Melville, 1957
Family Holasteridae Pictet, 1857
Genus Hemipneustes Agassiz, 1836
Hemipneustes compressus Noetling, 1897

Hemipneustes persicus Cotteau & Gauthier, 1895

Hemipneustes arabicus Ali, 1989

Order Spatangoida Claus, 1876
Family Hemiasteridae Clark, 1917
Genus Hemiaster Agassiz, in Agassiz & Desor, 1847
Hemiaster hattaensis Ali, 1989

Hemiaster paronai Checchia-Rispoli, 1921
Genus Mecaster Pomel, 1883

Mecaster victoris (Lambert, 1932)

Family Schizasteridae Lambert, 1905

Genus Linthia Desor, 1853

?Linthia sudanensis (Bather, 1904)
Genus Proraster Lambert, 1895

Proraster geayi Cottreau, 1908

124

Our field collecting in this region has more than doubled
the known fauna of late Cretaceous echinoids, making it one
of the most diverse assemblages of this age anywhere in the
world. Much of the material is well preserved, allowing
plating details to be recorded in many species and genera for
the first time. Forty four species are recognized (Table 1) and
sufficient material has been collected (over 2,500 specimens)
to allow a detailed biometric study of most species. Further-
more, bed by bed collecting has enabled me to recognize
recurrent assemblages and to place these into their palaeoen-
vironmental setting, something that has never previously
been attempted.

RELATIVE ABUNDANCES OF SPECIES

A total of 2585 specimens were collected and identified to
species level, allowing a quantitative assessment of species
abundance. This is not entirely accurate in comparison with a
previous study of echinoid abundance in the Cenomanian
(Smith 1988) where every specimen seen was collected,
because the three or four most abundant species are surely
undercollected. A few species (e.g. Petalobrissus rawdahensis
at Jebel Rawdah, section 2, or ‘Globator’ bleicheri at Jebel
Buhays) were so abundant that only a proportion of the
observed specimens were eventually collected. There is
therefore a bias towards the rarer species. Nevertheless, a
number of general conclusions can be drawn from the distri-
butional and abundance data that have been collected.

(a) There are approximately equal numbers of regular and
irregular echinoid species (22 regular, 23 irregular) (Fig. 1B).
A similar ratio of regular to irregular species is characteristic
of the present day biota (Kier 1977) and for extensive
collections from the Cenomanian of Great Britain and France
(Smith 1988, Neraudeau & Moreau 1989). Amongst the
irregular echinoids, cassiduloids (10 species) are more diverse
than holectypoids (5 species), spatangoids (5 species) or
holasteroids (3 species).

(b) Irregular echinoids greatly exceed regular echinoids in
numbers of specimens (Fig. 1B). In total 499 specimens of
regular echinoid were collected (19%), and this is likely to
overestimate their true relative abundance, since the five
most common echinoids were all irregular species and are
likely to have been significantly undercollected. This com-
pares well with previous studies, where Smith (1988) found
between 10-20% of the total number of echinoid specimens
collected (1800 individuals) from the Cenomanian of Wilm-
ington were regular species, and with Neraudeau & Moreau
(1989) who reported 22.8% of 5133 individuals collected from
the Cenomanian of North Aquitaine were regular species.
Therefore regular echinoids achieve a similar species diver-
sity to irregular echinoids, but occur in much lower absolute
diversity.

(c) Most species are rare. In terms of numbers of individu-
als, just five species make up more than 67% of the total
collection (Fig. 1A). Four of these five species are cassidu-
loid, the fifth (8% of the total) is the holectypoid *‘Globator’.
The most abundant species are cassiduloids and two of the
three species of Petalobrissus together form more than 36%
of the total number of individuals collected. In reality these
species are even more dominant, since they were certainly

A.B. SMITH

undercollected in comparison to rare species, possibly by as
much as a factor of two.

In comparison, the most abundant regular echinoid species
(Echinotiara perebaskinei and Glyphocyphus hattaensis) rep-
resent only 3.2% of the total collection each, and the the
most common spatangoid (Mecaster victori) and holasteroid
(Hemipneustes persicus) represent a mere 1.6% and 1.4%
respectively of the total collection.

ECHINOID PALAEOECOLOGY

Echinoids are adapted to live in a wide range of habitats and
the relationship between skeletal structure, habitat and mode
of life is now relatively well understood. Consequently they
are an excellent group to use as palaeoenvironmental indica-
tors. Virtually nothing is known about the late Cretaceous
palaeoecology of echinoids in tropical carbonate shelf envi-
ronments, and the Oman Mountains collection described
here provides the first such opportunity to assess echinoid
distribution and faunal associations quantitatively.

Before discussing the assemblages that can be recognized

and their palaeoenvironmental setting, it is necessary to look

at the detailed functional morphology of the different taxa.
Each has a preferred habit and life-style that is partially
reflected and can be deduced from the skeletal morphology
(e.g. Smith 1984).

1. Regular echinoids. All regular echinoids live epifaunally,
but can be differentiated into a number of ecological groups
based on their skeletal morphology.

(a) Hard-ground dwellers living within the zone of active
wave surge (0-5 m depth). This includes Codiopsis, Phy-
mechinus, Circopeltis and Echinotiara. These forms are char-
acterized by their flat oral surface and enlarged phyllodes
composed of P3/P4-type pore-pairs (Smith 1978). They also
typically have dense or modified aboral pore-pairs indicative
of specialized respiratory tube-feet. The broad, flat base and
numerous, large oral tube-feet are features characteristic of
present-day echinoids living on rocky surfaces within the zone
of active wave surge. Large and strong oral tube-feet are
necessary to provide adhesion in a turbulent environment.
Modern analogues would be Arbacia, Anthocidaris, Helioci-
daris or Stomopneustes, all of which are coastal species living
within the first few metres on rock or other hard ground
surfaces. They are rock grazers, feeding on filamentous or
fleshy algae which they obtain by rasping hard substrata.

(b) Hard-ground dwellers living subtidally within the top
few metres of water but subject to limited wave surge only.
Here I include only Goniopygus. Goniopygus has moderately
strong phyllodes, a depressed profile though without a broad
flat oral surface, and well-developed aboral respiratory tube-
feet (to judge from its pore-pairs). The aboral respiratory
tube-feet suggest it had a relatively high metabolic rate and
therefore lived in the shallow, warmest waters. It had strong
oral tube-feet for adhesion, but these were not as highly
developed as those of species in group (a) and it seems
unlikely that Goniopygus could have lived in fully exposed
habitats.

(c) Shallow water (ca. 2-10 m) forms living in more
protected environments, within wave base, but not subject to
strong currents or wave surge. These echinoids might typi-

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

Jebel Huwayyah, sections1/2:
50-4 Loftusia facies

404 N=57

Numbers of individuals

Faujasia

Species

100)

Jebel Huwayyah, section 1:
50-4 beds 14,15

40-4 N=62

Numbers of individuals

20
Species

Globator
bleicheri

Fig. 2 Echinoid species abundances within specific units at Jebel Huwayyah. A, fauna from beds 10-12, section 1 and beds 1-6, section 2
(Loftusia-rich facies). B, fauna from beds 14 and 15, section 1. Species are listed numerically along the x-axis as in Table 1. N = total

number of specimens collected.

cally be expected to have lived on or close to firm bottoms,
i.e. either rocky or reefal knolls or stabilized sedimentary
‘substrata. Taxa included here are Orthopsis, Phymosoma,
| Plistophyma, Actinophyma, Mimiosalenia, and Glyphop-
/neustes. These echinoids all have a depressed profile and
‘slight phyllodes, and often have increased densities of aboral
_pore-pairs, possibly specialized for gaseous exchange. Their
|spines are moderate to long. Modern equivalents, such as
Lytechinus or Temnopleurus live in and around hardground
substrata in reasonably well-protected habitats. They are
predominantly grazers, feeding on encrusting or boring algae
‘or plants and removing upright algae down to the substratum.
Those with no phyllode development and invaginated peris-
tomes are likely to have been grazers, whereas those with oral
phyllodes and a flat peristome were probably raspers.
Heterodiadema is a large, motile diadematoid-like echinoid
with long spines whose closest modern counterparts are forms
such as Centrostephanus or Diadema, found living in algal
turf (dense stands of filamentous algae) or in amongst thick-
ets in the reef flat environment (Birkeland 1989).

The cidarids Rhabdocidaris and Prionocidaris probably
delong to this category. They are clearly shallow-water forms
yecause of their well-developed respiratory tube-feet (conju-
a pore-pairs), but are globular in profile and lack phyl-
odes. They would have been confined to the most protected
f the shallow-water habitats, along with Heterodiadema.
‘idarids and diadematoids are rather generalist feeders,
referentially grazing on animals and plants, but also able to
ake up bottom material (Birkeland 1989).

(d) Protected subtidal (10 m plus) soft-bottom substrata

below active wave base. Noetlingaster, Hattopsis and Salenia
are all globular forms lacking aboral specialised respiratory
tube-feet. Noetlingaster closely resembles the present-day
Echinus or Tripneustes in morphology and presumably lived
in much the same way, found largely on sandy substrata of
lagoons or below active wave-base moving over stable sedi-
mentary bottoms or living within algal stands. These forms
are predominantly herbivorous browsers, cropping exposed
algae and taking loose bottom material. The extreme globu-
lar shape of Hattopsis is matched by that seen in some present
day temnopleurids such as Microcyphus and, like those
echinoids, it may have lived within algal turf, attached to and
enveloped by fronds of filamentous algae.

Irregular echinoids. These too can be divided into a number
of discrete ecological groups based on observed skeletal
characteristics.

(e) Infaunal medium-fine sand-grade burrowers which are
selective deposit feeders using penicillate tube-feet around
the mouth to pick up food particles from sediment of the
burrow floor. Here I include Hemiaster and Mecaster. Both
have globular tests with no real frontal sulcus. Pore differen-
tiation shows that funnel-building tube-feet would have been
present in ambulacrum III adapically, and subanally in the
posterior ambulacra also. These heart urchins clearly lived
infaunally within relatively poorly permeable, rather fine-
grained sediment. An apical funnel is required by infaunal
spatangoids living in finer-grained sediments, as is the aboral
fasciole (which is essential for maintaining a water-filled
space surrounding the test). The lack of specialization of the

126

100 Jebel Rawdah, section 1:

beds 3-5

N=170

Hattopsis
sphericus

Numbers of individuals

A.B. SMITH

, Globator
bleicheri

, Petalobrissus

Species

Fig. 3 Echinoid species abundances within beds 3-5 at Jebel Rawdah, section 1. Species are listed numerically along the x-axis as in Table 1.
N = total number of specimens collected. Unshaded box = fragments only.

frontal groove implies that all food particles were derived
from within the sediment, not from the sediment/water
interface.

(f) Infaunal medium-fine sand-grade burrowers, selective
deposit feeders harvesting organic material from the
sediment-water interface. Only Proraster falls into this group.
It has a deep, highly specialized frontal groove which is
characteristic of those modern taxa feeding by means of a
mucus-string. It is very similar to the extant Schizaster, and
presumably lived in very much the same way. The pore-pairs
in ambulacrum III are specialized and indicate the presence
of highly developed funnel-building tube-feet. There is little
doubt that Proraster lived infaunally, feeding on detritus from
the water/sediment interface that cascaded down the funnel.
The implication (though untested as far as I am aware) is that
Proraster is adapted for life in more organic-poor sediments
than either Hemiaster or Mecaster since it preferentially
harvests surface detritus. The well-developed petals show
that Proraster was still a relatively shallow-water form.

(g) Shallow infaunal or semi-infaunal ploughers in stable,
unconsolidated bottoms. Bulk deposit feeders harvesting
sediment at or close to the water interface. Only Hemip-
neustes falls into this catagory. Hemipneustes has a well-
developed anterior groove with specialized grill spines and
tube-feet. Sediment would have entered the frontal groove
adapically and passed to the mouth via a mucous string
running down ambulacrum III. The lack of protection for
petal tube-feet, and the asymmetry of the petals implies that
the surrounding sediment was highly permeable and that the
urchins lived only relatively shallowly burried.

(h) Mobile high permeability (low fines) unconsolidated

medium-coarse sands: infaunal bulk sediment swallowers.
This category includes Petalobrissus, Nucleopygus, Pyguros-
toma and Stigmatopygus. These urchins most closely
resemble the present day Apatopygus, which is a bulk sedi-
ment swallower. They have small phyllodes around the
peristome, with moderately well developed bourrelets (spe-
cialized regions of dense spines used for manipulating sand-
grade particles into the peristome. Their large periproct and
anal sulcus suggests they had to cope with copious faecal
discharge. They are small, depressed forms streamlined for
moving through loose sediment and have well-developed
petals.

Vologesia and Arnaudaster have similar phyllodes but |
much larger animals, more closely resembling Echinolampas.
Echinolampas is found today in subtidal (5-20 m) depths
living infaunally in shell-sands (e.g. Thum & Allen 1975).
Petalobrissus is likely to be adapted for more mobile sedi-
ments, i.e. for shallow subtidal shoals.

(i) Mobile, permeable unconsolidated sands: selective,
infaunal deposit feeders. Here I place Zuffardia and Fauyasia.
Both have small peristomes, surrounded by moderately well-
developed phyllodes, and with slight bourrelet development.
They both have rather small, posteriorly placed periprocts,
and from this it would appear that they were rather more
selective deposit feeders than Petalobrissus and other related
taxa. They have large petals and clearly lived infaunally,
because of their rounded shape. However, they could only
have done so within highly permeable sands, finer sediments
being insufficiently porous to allow sufficient water flow past
the well-developed petals.

(j) Infaunal selective particle feeders living within coarse,

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

A Mo Jebel Rawdah, section 2:

= Ad
on beds 3-10 5
2
90 N= 395 Dj i
% o
Yes
in c 8
2 so a
o 6
=) ies) <
3 Sh a
S 70 oo =
5 Ww Q eo)
£ >
—
Io}
ie
rs
E 40
2
30
20
10
Species
B a
700 2B
Jebel Rawdah, section 2: 2§
600-] beds 11, 12 Rs
oO
2 Bs
5004 N = 790 9 ss
Q
400 = g
5 3.9
GS YG
S 25
300 be ss
200 > oy
2 ao
100
90
2
& =
io} iy
S 70 i
2 3
S iS
i= a
6 5
g S
re 40
=|
2 0
oa)

Species

Jebel Rawdah, section 2:
200— beds 14-26

Petalobrissus
‘awdahensis

1804 N = 508
160
140
oO
120 3
Q
s
100 3
.s)
90 o
Ny
wo
7) a o
3% a = 3
7 o
z Ss 2 5 3
> 70 re) iS SS
so} 3 5 Ee
[= Q ‘= mB oS
re 2 = By no
S 8 2 Se alhitiuast
yo 0 a aS) o 2S
3 “ : eet ee
~-0 a i 2 a
wo ie)
= gs 8 38 5 &
= 90 € Qa is =
& 3 Y =
8 ro) a
2» st
2

Species

Fig. 4 Echinoid species abundances within units at Jebel Rawdah, section 2. A, fauna from beds 3-10; B, fauna from beds 11 and 12; C,
fauna from beds 14~26. Species are listed numerically along the x-axis as in Table 1. N = total number of specimens collected.

128 A.B. SMITH

Table 2 Inferred palaeoenvironmental settings for the late Cretaceous succession of the Oman-United Arab Emirates border region. The
common echinoid taxa found in each habitat are listed, together with indications of their probable life style.

Habitat A nearshore hardground dwellers living in zone of active wave surge.
(i) Hardground grazers: Echinotiara perebaskinei, Phymechinus? perplexus, Codiopsis lehmannae.

Habitat B nearshore hardground dwellers living in more protected environments (perireefal).
(i) Hardground grazers: Goniopygus arabicus.

Habitat C subtidal protected environments within normal wave base; mixed cobble and sand substrata with nearby level-bottom reefal thickets.
(i) Epifaunal grazers: Glyphopneustes hattaensis, Mimiosalenia quinquetuberculata, Goniopygus arabicus, Phymosoma hexoaporum.
(ii) Epifaunal browsers: Circopeltis? emiratus, Heterodiadema buhaysensis, Orthopsis miliaris, Hattopsis paucituberculatus, Plistophyma |

asiaticum.

(iii) Epifaunal generalists: Prionocidaris morgani, Prionocidaris? emiratus, Salenia nutrix, Salenia microprocta.

(iv) Infaunal grazers/detritivores: ‘Globator’ bleicheri, Conulus douvillei, Coenholectypus cf. baluchistanensis.
(v) Infaunal bulk sediment swallowers: Nucleopygus magnus, Pygurostoma morgant. }

(vi) Infaunal selective deposit feeders: Vologesia rawdahensis, Arnaudaster cylindriformis, Hemiaster hattaensis.

Habitat D Shore-face sand flats at or within normal wave base.
(i) Bulk sediment swallowers: Petalobrissus rawdahensis, P. ct. setifensis.
(ii) Selective deposit feeders: Zuffardia morgani, Faujasia eccentripora.
(iii) Infaunal generalist detritivores: ‘Globator’ bleicheri, Coenholectypus cf. baluchistanensis.

Habitat E Broad shallow, open shelf subtidal sand flats, within normal wave base (2-10 m) but distant from the platform margin and thus

relatively protected. i
(i) Epifaunal grazers: Noetlingaster paucituberculatus, Hattopsis sphericus.
(ii) Semi-infaunal selective deposit feeders: Hemipneustes spp.

(iii) Infaunal selective deposit feeders: Mecaster victoris, Hemiaster paronai, Stigmatopygus? pulchellus, Faujasia eccentripora.

(iv) Infaunal bulk sediment swallowers: Petalobrissus rawdahensis, P. cf. setifensis, Pygurostoma morgani.

Habitat F Deeper-water (20 m plus) platform basinal muddy sands, below normal wave base.
(i) Epifaunal grazers: possibly Actinophyma spectabile.
(ii) Infaunal selective deposit feeders, harvesting detritus from the sediment/water interface: Proraster geayi.

permeable sands behind fringing reefs in 0-10 m water depth. stabilized bottoms in shallow water conditions protected from
Conulus, ‘Globator’ and Coenholectypus belong here, though the influence of strong wave action. The presence of ‘Globa-
the highly inflated C. inflatus may have reverted to a prima- tor implies that the sediment was unconsolidated, permeable
rily epifaunal mode of life, below normal wave base. Globa- and rather coarse, while horizons with an abundance of |
tor closely resembles the present day Echinoneus whose Hattopsis might suggest the presence of nearby algal stands.

ecology was described by Rose (1976). Echinoneus lives The abundance and diversity of regular echinoids (most of
beneath coral debris in coarse shell-sands passing suitable which are algivore grazers) implies a mixture of environments |

grains into its peristome by means of its tube-feet. were close by, ranging from rocky (?patch-reef, as seen at
Inferred depths and habitats for typical Omani echinoids Jebel Faiyah) to stabilised sedimentary bottoms with algal |
are tabulated in table 2. stands. The palaeoenvironment is therefore most likely to

represent a shallow (5-10 m) backreef or leaward environ-
ment that supports a high algal diversity.

ECHINOID ASSEMBLAGES (2). Echinotiara/Faujasia/Phymechinus assemblage. An
assemblage which includes a number of rarer regular echi-
Because careful count of the number of specimens collected noids absent from most other assemblages (e.g. Codiopsis,
for each taxon was kept it is possible to identify recurring Phymechinus). Faujasia is the only common element found at
assemblages. The following seven assemblages are differenti- other levels, but no other irregular echinoid occurs in any |
ated here. abundance (Fig. 4A).
This assemblage is best developed at Jebel Rawdah section
(1). Conulid/arbacioid assemblage. An abundance of Conu- 2, in the basal calcarenitic beds with associated coral/rudist
lus and ‘Globator’ which is usually accompanied by a diverse debris.
mixture of regular echinoids, notably by the arbacioids Interpretation: this assemblage is dominated by hard-
Goniopygus, Glyphopneustes, Mimiosalenia and Hattopsis. ground, shallow-water (0-5 m) epifaunal regular echinoids
Other taxa making up a smaller component of the fauna (Echinotiara, Phymotaxis, Codiopsis) adapted for life in
include Phymosoma, Nucleopygus and Pygurostoma (Figs strong wave-surge environments. Clearly none are preserved
2B, 3, 5B, 6). in situ, but have been transported into immediately adjacent
This assemblage is characteristic of the lower beds of the basins of sediment accumulation, along with coral and stro-
Simsima Formation at Jebel Buhays, Jebel Thanais, Jebel matoporoid debris and hippuritid rudist debris indicative of
Faiyah, and Jebel Huwayyah. It is typically developed with reef habitat. Transportation has not been far, otherwise much
the income of relatively coarse, carbonate bioclastic sands in more disintegration of tests might be expected. The occur-
the succession. rence of large blocks of rolled coral with stromatoporoid and

_Interpretation: the conulid/arbaciid assemblage has a very rudist debris at Jebel Rawdah section 3, implies that the
high diversity of regular echinoids that would have preferred deposits accumulated close to actual reefal framework. Fauja-

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

, Mecaster victoris

Petalobrissus
rawdahensis

Globator
" bleicheri

Species

A *°] Jebel Rawdah, section 3: beds 1-4 _E@
g 50 N = 88

g Jebel Rawdah, section 4: beds 1-7
ao N=25

w 30

5 20

5 10

z ee

| Jebel Rawdah, section 3: beds S+
@ sof N= 53

zg Jebel Rawdah, section 4: beds 8+ §S
> 1 N=49

w 30

5 20

5 10

60 : ;
| i Jebel Faiyah, section 1: beds 8-10
! —_
w =
S 59 N=77
3
| 2 40 s
aS) o
| £ a 3
—
fo} a)
n es
@ 20 28
| E 10 es
| z Ss
5 10 15 20

Table 1. N = total number of specimens collected.

fa may be the only in situ echinoid for the depositional
avironment, living within the surrounding sand-fields in an
nmediately subtidal environment, although none is pre-
‘rved with associated spines. Goniopygus is found in asso-
ation, and it probably lived amongst the tallus derived from
ie reef crest rather than on the crest itself.

) Petalobrissus assemblage. At certain levels of coarse,
consolidated orbitoline limestones Petalobrissus occurs in
ige abundance, accompanied by much smaller numbers of
her infaunal echinoids (Zuffardia, Faujasia, Coenholecty-
js and ‘Globator’). Rare Noetlingaster are the only regular
hinoids encountered (Fig. 4B).

Globator
bleicheri

. Species

)Fig.5 Echinoid species abundances within units at Jebel Rawdah, sections 3 and 4. A, fauna from lower beds; B, fauna from higher beds.
Species are listed numerically along the x-axis as in Table 1. N = total number of specimens collected.

Globator
bleicheri

Species

‘ig.6 Echinoid species abundances within units at Jebel Faiyah, section 1, beds 8-10. Species are listed numerically along the x-axis as in

This assemblage is seen only in the lower part of the section
(beds 11-13) at Jebel Rawdah, section 2.

Interpretation: this assemblage is composed entirely of
small, bulk sediment swallowers and is suggestive of shallow
subtidal unconsolidated and well-washed sand flats free of
algae, presumably subject to constant disturbance. The beds
can thus be envisaged as shallow (0-10 m) orbitoline sand
flats developed behind a fringing reef. The abundance of
individuals suggests that this was a region of relatively high
productivity.

(4) Hemipneustes/Noetlingaster assemblage. The abundance
of large infaunal selective deposit feeders, notably Hemip-

130

neustes and Pygurostoma, characterises this assemblage. Usu-
ally Noetlingaster-rich levels alternate with holasterid-rich
levels, so it may not be entirely correct to treat these as one
assemblage, though they occur intermixed within the same
lithofacies. All the species are large in comparison to those
elsewhere. This assemblage typifies the upper beds of section
2, Jebel Rawdah (Fig. 4C).

Interpretation: the Hemipneustes/Noetlingaster assemblage
is composed of selective deposit feeders specialized for life
within coarse, permeable sands, together with occasional
specific bands of the large generalist algivore Noetlingaster.
The dominance of large selective deposit feeders implies a
relatively nutrient-starved environment where large-scale
harvesting of the sediment/water interface is needed to
exploit the food supply. Associated lithofacies evidence sug-
gests that they were living within shallow-water (2-10 m)
flat-bottom shoals, probably immediately infratidal. Noetlin-
gaster, like Echinus in today’s seas, may have been a general-
ist detritivore/algivorous browser. The environment of
deposition is therefore envisaged as relatively quiet sedimen-
tary level bottom shoals that lay not particularly deep but
distant from the high productivity reef area or from more
exposed platform margin conditions.

(5) Faujasia assemblage. Occasional, almost monospecific
assemblages of Faujasia are found in calcified fine sands at
Jebel Huwayyah near the base. In the associated Loftusia-
rich levels there are also occasional Hemiaster and Proraster.
These may be nearshore infratidal sands in relatively pro-
tected environments (Fig. 2A).

(6) Mecaster assemblage (Fig. 5A). Another virtually
monospecific assemblage confined to a single level at Jebel
Rawdah section 3. This time the only associated echinoids are
the rare Hemipneustes.

(7) Proraster assemblage. Only Proraster is found in the
highest, muddiest limestones of Jebel Huwayyah. The rela-
tively fine-grade sediments and the absence of infaunal
cassiduloids or holasteroids suggests that these beds may
represent shelf-basinal deposits more than ca. 20 m deep.
Only highly specialist infaunal forms such as Proraster could
apparently cope with life in these relatively nutrient-poor
mud-rich deposits.

PALAEOECOLOGICAL SYNTHESIS

The faunal succession records a relatively rapid deepening
from beach boulder beds and sands through nearshore,
high-energy, subtidal conditions with off-beach coral-rudist
reefal patches to somewhat deeper-water conditions at or
immediately beneath wave base. Maximum echinoid diversity
is found in the shallow sand fields strewn with shell and coral
debris surrounding coral thickets that formed at about wave-
base around the shores of the uplifted ophiolite. Subsequent
shallowing over the platform led to the creation of broad,

shallow shoals supporting a low-diversity infaunal echinoid |

assemblage, dominated by holasteroids and cassiduloids. The
only epifaunal echinoid here is Noetlingaster. The deepest
water facies are probably those seen at the top of the section
at Jebel Huwayyah, where only the infaunal spatangoid
Proraster occurs.

SYSTEMATIC DESCRIPTIONS

Class ECHINOIDEA Leske, 1778
Subclass CIDAROIDEA Claus, 1880
Order CIDAROIDA Claus, 1880
Family RHABDOCIDARIDAE Lambert, 1900

Gen. et sp. indet. Pl. 1, fig. 1; Figeey

MATERIAL. A single specimen, comprising three interambu- |

lacral plates, BMNH EE3438.

OCCURRENCE. From the scree at Jebel Buhays, section 1;
derived from the lowest 2-3 metres of the Simsima Forma-
tion.

DESCRIPTION. Three interambulacral plates, rather badly
weathered, from the ambital region of the test. Plates are,
18-7 mm wide by 8 mm tall and appear to have tesselate
sutures. Each plate carries a single large tubercle with a
circular, confluent areole that occupies the full height of the
plate. The boss is surmounted by a broad, strongly crenulate

Lithofacies 4

Well-sorted orbitoid-

rich packstones oe
|
!
|

Hemipneustes spp.
Coenholectypus inflatus Tithotacies) 3b
Pygurostoma
Stigmatopygus

3 Poorly-sorted orbitoid-
Noetlingaster y

rich packstones with
thodolites and Dictyoptychus

Petalobrissus spp.

Zuflardia
Hattopsis
Hemiaster

Lithofacies 3c |

Muddy calcarenites: very
poorly sorted. Loftusia and
Cunnolites prominent.

I Bedrock

| | Shore-face

I Echinotiara | clastic sands
Codiopsis

Phymechinus? Faujasia

Lithofacies 3a

Poorly-sorted bioclastic
calcarenites

Goniopygus

Glyphopneustes

Salenia

Orthopsis

Phymosoma

Nucleopygus

Globator

Coenholectypus baluchistanensis

}

Fig. 7 Schematic reconstruction of the sea-floor in Maastrichtian times showing the range of environments represented in the study area. See
urchins are illustrated in their probable mode of life and the inferred environmental ranges for key taxa are shown.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

Fig. 8 Camera lucida drawing of an interambulacral plate of
Rhabdocidaridae gen. et sp. indet., BMNH EE3438. Scale bar =
5mm.

platform while the mamelon itself is rather small (1-2 mm
diameter) and has a large central perforation. The primary
tubercle lies subcentrally on the plate with a broad adradial
and interradial platform on either side. The broad zones
outside the areole of the primary tubercle have a few scat-
tered secondary tubercles preserved, but are otherwise too
abraded to retain any evidence of fine tuberculation that may
have been present. It is clear from the distribution of the
tubercles that are preserved that these regions were covered
in a rather heterogenous and open array of various-sized
tubercles.

REMARKS. The broad plates, confluent areoles, perforate
crenulate tubercles and heterogeneous secondary tubercula-
tion show this to be a member of the Rhabdocidaridae. It is
impossible to place this specimen in any genus on the basis of
such incomplete material. However, it most probably repre-
sents a species of Rhabdocidaris itself, judging from the size
of the specimen and the coarseness of the secondary tubercu-
lation.

Family CIDARIDAE Gray, 1825
Tribe CIDARINI Gray, 1825
Subtribe PHYLLACANTHINA Smith & Wright, 1989
Genus PRIONOCIDARIS Agassiz, 1863

Prionocidaris morgani (Gauthier, 1902) Pl. 1, figs 2-4;
Fig. 9B

Rhabdocidaris (Leiocidaris) morgani Gauthier: 145,
pl. 20, figs 3-6.

1989 Cidaris cf. scabra Gauthier; Ali: 398, fig. 2.1.
_ Types. The syntypes are the two specimens described and
figured by Gauthier. One is a more or less complete test 35
_mm in diameter, the other an interambulacral segment. The
whereabouts of this material is unknown: the specimens are

1902

131

not in the Morgan collection in the Museum National
d'Histoire Naturelle, Paris.

MATERIAL STUDIED. Two specimens: BMNH EE3433, from
the scree derived from the lowest beds of the Simsima
Formation at Jebel Buhays, section 1; BMNH EE34335, loose
in the scree at section 3b, Jebel Rawdah. A third fragment,
EE3434, probably belonging to this species, was also found
loose in the scree at Jebel Buhays.

OCCURRENCE. The syntypes come from the late Cretaceous
(‘Senonien superieur’) of Louristan, Iran (Gauthier was
unable to give a precise locality). In the United Arab
Emirates the species comes from the Maastrichtian of Jebel
Buhays and Jebel Rawdah.

DIAGNOSIS. A species of Prionocidaris with broad conjugate
pore-pairs occupying more than half of the ambulacral plate
width, and a perradial zone with a single primary tubercle and
a row of tubercle-like granules, up to four abreast, on each
plate. Interambulacra with well differentiated scrobicular
circles and aligned extrascrobicular tuberculation.

DESCRIPTION. The following description is based on BMNH
EE3433, the larger and better preserved of the two. The
specimen comprises parts of two interambulacra and ambu-
lacra. Interambulacral width at the ambitus is 23 mm, sug-
gesting a test diameter of approximately 50 mm in life. Test
height is 32 mm. The ambulacra are 6 mm in width and only
slightly sinuous. The pore-pairs are very wide and conjugate
(Pl. 1, fig. 4; Fig. 9B) and successive pore-pairs are closely
spaced. The poriferous zone occupies approximately half of
the plate width. There is a single primary tubercle with a
mamelon immediately adjacent to each pore-pair. The
remainder of the perradial zone is occupied by a row of large
granules (non-mamelonate), some three or four abreast and
organised into discrete vertical rows.

Interambulacral plates are relatively broad, an ambital
plate being 13-1 mm by 9-4 mm in height. All plates, except
the most adapical in each column, have a single large primary
tubercle. This has a sunken areole 7-2 mm in diameter and a
mamelon 2.2 mm in diameter (PI. 1, figs 2, 4). Mamelons are
perforate and the surrounding platform is non-crenulate. The
primary tubercles lie offset towards the adradial suture leav-
ing a broad interradial zone of miliary tuberculation. There is
a clearly differentiated ring of scrobicular tubercles surround-
ing each areole. Extrascrobicular tuberculation is well devel-
oped with approximately 2 miliary granules abreast towards
the adradial suture, eight abreast towards the interradial
suture, and either a single row or no row developed adapical
and adoral to the scrobicular circles. These extra-scrobicular
tubercles are non-mamelonate and decrease in size towards

Fig.9 Camera lucida drawings of ambital ambulacral plating in cidarid species. A, Stereocidaris persica (Cotteau & Gauthier), Museum
@ Histoire Naturelle, Morgan Collection; B, Prionocidaris morgani (Gauthier), BMNH EE3433; C, Prionocidaris? emiratus sp. nov.,

|
|
| BMNH EE3431. Scale bar = 1 mm.

o>)
bo

A.B. SMITH

PLATE 1
Fig. 1 Rhabdocidarid gen. et sp. indet. BMNH EE3438; three interambulacral plates, x 3. Jebel Buhays, section 1; loose in the scree

derived from the lowest 3 m of the Simsima Formation.

Figs 2,4 Prionocidaris morgani (Gauthier). BMNH EE3433. 2, adapical portion of interambulacrum and adjacent ambulacral zones, X 2:5;
4, interambulacrum detail, x 4-5. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation.

Figs 3, 5,6 Prionocidaris? emiratus sp. nov. BMNH EE3431, holotype. 3, ambulacral zone detail, x 10; 5, lateral, x 2; 6, oral, x 2. Jebel
Huwayyah, section 2, beds 3-6.

Fig. 7 Cidarid spine, possibly belonging to Prionocidaris morgani (Gauthier). BMNH EE3434, x 3. Jebel Buhays, section 1; loose in the
scree derived from the lowest 3 m of the Simsima Formation.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

the interradius. There are neural grooves developed in the
extra-scrobicular regions.

The smaller specimen (BMNH EE3435) is more complete,
but less well preserved. It has seven interambulacral plates in
a column at an estimated test diameter of about 35 mm.
Adapical interambulacral plates retain fully formed tubercles.

Two cidarid spines were also collected from the scree at
Jebel Buhays and probably belong to this species. The best
preserved is BMNH EE3434 (PI. 1, fig. 7). It is 19 mm long
by 2-6 mm wide. It has a short neck and a perforate,
non-crenulate base. The shaft is widest midlength and tapers
towards the tip, but then expands slightly at the very tip to
end in a blunt crown. The shaft is covered in rows of thorns
which coalesce to form ribs towards the tip.

REMARKS. This species differs from the most common late
Cretaceous cidarid described from Iran, ‘Cidaris’ persica
Cotteau & Gauthier (1895), in having much finer and denser
extra-scrobicular tuberculation and wider and more strongly
conjugate pore-pairs at a comparable size (compare Figs 9A,
B). ‘C’. persica belongs to the genus Stereocidaris and has a
characteristic strong interporal ridge developed between
pore-pairs. Cidaris scabra Gauthier (1902) was established
for a 30 mm diameter individual of uncertain provenance
within Louristan. It differs primarily in having two ambulac-
ral tubercles to each plate but has very similar strongly
conjugate pore-pairs. Ali (1989) described a specimen of P.
morgani from Jebel Rawdah under the name Cidaris cf.
scabra Gauthier.
Spines comparable in morphology to the one described
here have previously been described under the name Cidaris
aftabensis Gauthier (1902). They can only tentatively be
assigned to this species until specimens are found that are
attached to a test.

-Prionocidaris? emiratus sp.nov. Pi. figs 3, 5,6; Fig.
XC

TyPE. Holotype and only known specimen, BMNH EE3431.

) OCCURRENCE. Found loose in the Loftusia levels a little
\ below Beds 6/7, section 2, Jebel Huwayyah.

‘DIAGNOs!s. A cidarid with conjugate ambulacral pore-pairs
and a single primary ambulacral tubercle to each plate,
adjacent to the pore-pair. The perradial zone of each ambu-
lacrum is filled with small, dense miliary tuberculation, three
or four to a plate and arranged in two rows.

DESCRIPTION. Test 28-5 mm in diameter by 18-9 mm in
height. Ambulacra 3-6 mm in width at the ambitus, slightly
sinuous. Pore-pairs strongly conjugate, with individual pores
af each pair ca. 0-3 mm diameter and separated by an
|nterporal distance of about 0-7 mm (PI. 1, fig. 3; Fig. 9C).
?erradial zone elevated and more or less flat; comparatively
‘hort and tall. A large primary tubercle occupies about half of
he tuberculate region on each plate (PI. 1, fig. 5; Fig. 9C).
These tubercles are almost contiguous vertically. The central
art of the perradial zone is occupied by very much smaller
tanules, some three or four to a plate, irregularly scattered.
| There are six interambulacral plates in a column, of which
1e most adapical in each zone has only a rudimentary
dbercle. Ambital plates are relatively tall and narrow, 7-0
im wide and 5-8 mm high (Pl. 1, fig. 5). The primary
ubercle lies centrally on the plate and has a sunken areole 4
1m in diameter (on ambital plates). The mamelon (1-1 mm

133

in diameter) is perforate and non-crenulate. The surrounding
scrobicular circles are non-contiguous. Extra-scrobicular
tuberculation is very fine and dense. There are approximately
three to four granules abreast on the interradial margin, three
abreast adapically and two abreast both adradially and
adorally.

REMARKS. The strongly conjugate pore-pairs indicate that
this is a member of the Phyllacanthina, and in many respects
it is close to P. morgani in tuberculation and appearance. The
interambulacra are narrower than usual for Prionocidaris,
and the interradial zones less well developed. Therefore, it is
only tentatively assigned to this genus. P? emiratus can be
easily differentiated from P. morgani by its ambulacral tuber-
culation (compare Figs 9B and 9C). None of the cidarid
species described by Cotteau & Gauthier (1895) or Gauthier
(1902) have this form of tuberculation.

Indet. cidarid plates

In addition to the specimens described above, fragmentary
material of cidaroids has been collected from the basal beds
at Jebel Faiyah, section 1, and the Loftusia levels (beds 1-6)
at Jebel Huwayyah. None of this material is adequate to
determine to generic level.

?Order DIADEMATOIDA Duncan, 1889
Family HETERODIADEMATIDAE Smith & Wright, 1993
Genus HETERODIADEMA Cotteau, 1864
Heterodiadema buhaysensis sp. nov. Pl. 2, figs 1-3;
Figs 10, 11

Types. Holotype BMNH

EE3442-5, EES019.
OTHER MATERIAL. One other specimen, BMNH EE3446.

BMNH EE3441; paratypes,

OCCURRENCE. Five specimens come from the lowest beds of
the Simsima Formation and were collected in the scree
derived from the lowest few metres of that formation at Jebel
Buhays, section 1. One was collected from the lowest 1-5 m of
Simsima Formation at Jebel Thanais. One specimen was
collected from bed 5 at Jebel Rawdah, section 3b.

DIAGNOsIS. Apical disc large, pentagonal, caducous; prob-
ably monocyclic to judge from the outline. Ambulacra
trigeminate, relatively wide with diadematoid-style plate
compounding. Phyllodes absent. Primary tubercles perforate,
crenulate; reducing in size sharply adapically and tending to
become imperforate. Broad zones of granulation along the
perradius, adradius and interradius. Peristome invaginated
with reasonably deep and sharply defined buccal notches with
tags.

DESCRIPTION. Tests range in diameter from 18-36 mm. The
test is circular in outline and depressed in profile, with a
height 42-48% of the diameter (mean = 45%; Fig. 10). The
ambitus lies at about mid-height and the sides are uniformly
rounded (PI. 2, fig. 3). The apical disc and peristome are
slightly invaginated in the largest specimen. Coronal plates
are all firmly sutured together.

The apical disc is caducous and has been lost from all
specimens. The apical disc opening is pentagonal in outline
with the angles projecting reasonably strongly into the inter-
radii (slightly more into the posterior interradius, which isalso

134

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Fig. 10 Biometric data for Heterodiadema buhaysensis sp. nov.

Kig. 11 Camera lucida drawing of three ambital ambulacral plates
of Heterodiadema buhaysensis sp. nov., BMNH EE3444. Scale
bar = 1 mm.

A.B. SMITH

more rounded). It is a little longer than broad with a length
that is 36-43% of the test length.

Ambulacra are broad and nearly parallel-sided at the
ambitus but taper gradually both adapically and adorally. At
the ambitus they have a width that is 23-25% of the test
length. Pore-pairs are uniserial throughout, without any
widening whatsoever at the peristomial margin. Plates are
compound, with diadematoid-style triads in which the central
element is the largest and upper and lower elements both
taper towards the perradial and adradial suture (Fig. 11).
Each triad bears a primary tubercle and there is a well-
developed perradial band of miliary tuberculation that
expands adapically. At the ambitus in the largest specimen
perradial tubercles are about three abreast. Oral and ambital
tubercles are perforate and crenulate, but tubercles reduce
sharply in size above the ambitus, where they may become
imperforate and non-crenulate (some remain perforate). This
reduction in size occurs above about the seventh compound
plate. Larger tubercles have circular, non-confluent areoles.
At the ambitus pore-pairs are rather widely spaced and
separated by a single row of miliary tubercles. There are no
obvious sphaeridial pits around the peristome.

Interambulacra are about 1-3-1-5 times as wide as the
ambulacra. There are 15 plates in a column at 36 mm test
diameter. All plates have a single large primary tubercle that
is perforate and crenulate and is surrounded by a circular
areole. The mamelon is notable for its small size in compari-
son to the size of the areole. Areoles are non-confluent, being
separated for the most part by a single band of miliaries.
Tubercles decrease in size sharply towards the apex and may
become imperforate. There are broad zones of uniform
miliary tuberculation both adradially and interradially: about
six abreast adradially and three or four abreast interradially
at the ambitus in the largest specimen. Scrobicular tubercles
are not differentiated.

The peristome is relatively large, 28% of test diameter in a
36 mm diameter individual. It is strongly indented by the
buccal notches. There are broad, smooth tongue-like regions
running adradially from each buccal notch and extending up
to the fourth interambulacral plate.

REMARKS. This species is most closely related to the mid-
Cretaceous (Cenomanian) Heterodiadema lybica (Agassiz &
Desor). Both have a similar form of ambulacral compounding
and a very similar tuberculation style. Granular zones are
broad and well developed and in both there is a sharp
reduction in the size of primary tubercles above the ambitus,
with those adapically tending to become imperforate and
non-crenulate. Finally, both have the distinctive tubercle-free
tag to the buccal notches and uniserial pore-pairs throughout.
H. buhaysensis differs from H. lybica in having fewer reduced
tubercles, the primary tubercles becoming smaller consider-
ably closer to the apex in H. buhaysensis in individuals of
comparable size. It also differs in lacking the extreme prolon-
gation of the apical disc into the posterior interambulacrum
seen in medium to large specimens of H. lybica.
Heterodiadema was placed in its own family by Smith &
Wright (1993). Where this family fits into the higher tax-
onomy of echinoids, however, is much less certain. They may
be members of the aulodont group Diadematoida. This is
suggested by the prominent buccal tags and the very delicate
mamelons on the primary tubercles, which indicate that the
species had small, fragile spines. Alternatively they may be
stirodonts, early members of the Phymosomatoida (as

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

) ATE 2

lis 1-3 Heterodiadema buhaysensis sp. nov. BMNH EE3441, holotype; 1, oral; 2, apical; 3, lateral; all x 3. Jebel Buhays, section 1; loose
m the scree derived from the lowest 3 m of the Simsima Formation.

Is4,5 Orthopsis miliaris (d’ Archiac). Topotype specimen of Orthopsis morgani Cotteau & Gauthier, from the Morgan Collection,
Museum d’Histoire Naturelle, Paris, x 2. Senonian, Khianan, Iran.

136

Test height (mm)

30 40 50

Test diameter (mm)

@ Ambulacral pore-pairs
® \nterambulacral plates

Number in column

Test diameter (mm)

Fig. 12 Biometric data for Orthopsis miliaris (d’ Archiac).

defined by Smith & Wright, 1993). Diadematoids and phy-
mosomatoids are fundamentally differentiated on the struc-
ture of their spines and lantern apparatus, diadematoids
having hollow spines and grooved teeth, phymosomatoids
having solid spines and keeled teeth. In neither H. /ybica nor
in our new species are the spines or lantern known, thus its
higher taxonomic position must remain unresolved. Because
of its delicate tuberculation, it is tentatively assigned here to
the Diadematoida.

Infraclass ACROECHINOIDEA Smith, 1984
Plesion (Order) ORTHOPSIDA Mortensen, 1942
Family ORTHOPSIDAE Duncan, 1889
Genus ORTHOPSIS Cotteau, 1864

Orthopsis miliaris (d’ Archiac, 1835) PI. 2, figs 4, 5; Pl.
3, figs 1-9; Figs 12-14

1835 Cidarites miliaris @ Archiac: 179, pl. 11, fig. 8.

1895 Orthopsis morgani Cotteau & Gauthier: 87, pl. 14, figs
6-9.

1933. Orthopsis sanfilippoi Checchia-Rispoli: 6, pl. 1, figs
5-15.

1985 Orthopsis miliaris (d’ Archiac); Geys: 143, pl. 5, figs
8-10 (see also for full prior synonymy).

1989 Orthopsis morgani Cotteau & Gauthier; Ali: 401, fig.
A(T).

1991 Orthopsis miliaris (d Archiac); Smith & Bengtson: 30,

pl. 8B-F, text-fig. 23.

A.B. SMITH |

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MATERIAL STUDIED. Forty seven specimens of which the |
following 11 were used in the biometric analysis: BMNH
EE3720-21, E3723, EE3725-26, EE37/28, EE373iy
EE3733, EE3738, EE3740, EE3749 and EES018.

OCCURRENCE. In the western Oman mountains this species |

occurs at the following localities and horizons (numbers in

brackets are number of specimens):

Jebel Buhays, section 1: loose in scree derived from lowest |
few metres of the Simsima Formation (13)

Jebel Thanais: lowest 2 m of Simsima Formation (1). |

Jebel Huwayyah, section 2: Loftusia levels (beds 3-5; 1|
fragment).

Jebel Rawdah, section 1: beds 3 and 4, and loose (4).

Jebel Rawdah, section 3b: bed 2 (1); bed 3, 1m above base
(1), bed 8 (2); bed 9 (8); bed 10 (1); loose (3).

Jebel Rawdah section 4: bed 12 (5);top of bed 15 (1); loose, |
lower half of sandy beds (1).

Jebel Faiyah, section 1a: top of bed 2 (1).

DESCRIPTION. Tests range from 20 mm to 48 mm in diameter
and are circular in outline and bun-shaped in profile. Test
height is 39-52% of test diameter (Fig. 12) and in profile the
ambitus lies about one third the height above the base.

The apical disc is dicyclic, though occasionally one of the
posterior oculars may just be exsert. The apical disc occupies
17-24% of the test diameter (mean = 21%, SD = 1-:9%, N=
10; Fig. 12). Genital plates are broad and crescentic in
outline, except for the madreporite, which is larger and more
pentagonal in outline (Pl. 3, fig. 7; Fig. 13A). Madrepores
occupy almost the entire surface of the madreporite plate and

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

there are small scattered tubercles amongst the openings.
Gonopores are present even in the 20 mm diameter indi-
vidual. Ocular plates are small and pentagonal. All plates
have small secondary tubercles, those on the genital plates
tending to form a circle around the periproct. The periproct is
irregularly oval in outline and occupies 10-14% of the test
diameter.

Ambulacra are 17-21% of the test diameter in width at the
ambitus (mean = 20%, SD = 1:3%, N = 11). Plating is
trigeminate throughout and pore-pairs are arranged either

Fig. 13. Camera lucida drawings of plating in Orthopsis miliaris
(d’Archiac). A, apical disc plating, BMNH EE3723; B, ambital
ambulacral (left) and interambulacral (right) plates, BMNH
EE3723; C, adoral plating, ambulacrum to left, interambulacrum
to right, BMNH EE3733. Scale bars: A, B = 1 mm; C = 2 mm.

Fig. 14 Camera lucida drawings of lantern elements in Orthopsis
miliaris (d’ Archiac): A-D, BMNH EE3735; E, BMNH EES5020.
A, pyramid with rotula and tooth in position; B, same in lateral
view; C, fragment of compass element; D, adapical view of
pyramid, with proximal end of tooth (broken) towards the top; E,
cross-section of a single tooth. Scale bars = 1 mm.

137

uniserially or in very weak arcs of three (PI. 3, fig. 6). All
ambulacral elements are narrow and elongate and reach the
perradius (Fig. 13B). A primary tubercle (perforate and
non-crenulate) straddles two of the three elements in each
compound plate (PI. 3, fig. 6). The third element carries two
small secondary tubercles and an intermediate row of miliary
granules. There are secondary and miliary tubercles down the
perradius also. Adorally only the first five or so pore-pairs are
offset to form a weak phyllode (Fig. 13C). There are 55
pore-pairs in a column at 20 mm test diameter, rising to
around 90 at 46-48 mm diameter (Fig. 12). Sphaeridial pits
are very shallow and are present on the two or three most
adoral compound plates, immediately adjacent (perradial) to
the pore-pair on the element in each triad that does not
support a primary tubercle.

Interambulacra are 38-44% of the test diameter in width at
the ambitus (mean = 41%, SD = 2:0%, N = 11) (Fig. 12).
Ambital plates are short and wide and slightly curved. At the
ambitus most specimens have a single primary tubercle,
centrally positioned, plus two smaller secondary tubercles
one on either side (PI. 3, figs 3, 8; Fig. 13B). However, in a
fragment of a large specimen (ca. 65 mm diameter, BMNH
EE3717), there is a fourth large tubercle on the interradial
margin. The remainder of the plate carries scattered tertiary
tubercles and granules. All tubercles are perforate and non-
crenulate. Adorally there is no primordial plate. There are 14
plates in a column at 20 mm test diameter, rising to 18 at
42-48 mm test diameter (Fig. 12).

The peristome is 36-40% of the test diameter in diameter
(mean = 38%, SD = 1:7%, N = 7). It is slightly invaginated.
There are strong buccal notches that extend into the second
interambulacral plate (PI. 3, fig. 9; Fig. 13C).

The perignathic girdle is seen in BMNH EE3733. The
auricles are well developed. They meet and are fused above
the perradius to form a continuous arch.

Lantern elements are preserved within the tests of several
specimens, but are best seen in BMNH EE3735. Hemipyra-
mids are tall and have a relatively shallow foramen magnum,
only some 33% of their height (Figs 14A, B). There is a clear
muscle attachment flange on the outer edge of each. The
upper surface of the lantern is also pitted. There are well-
developed processus superalveolares that extend to the tooth.
The lateral (inter-pyramidal) face is denticulate along its
inner edge and has the usual horizontal series of ridges for
muscle attachment.

The epiphyses are not clearly seen in any specimen but
appear relatively short. They are definitely not extended
above the foramen magnum nor are they fused together as is
the case in camarodont lanterns. There is a single compass
element that is siender and has a small head (Fig. 14C). The
tooth is strongly keeled and is best seen in cross section in
BMNH EE5S020 (Fig. 14E).

Spines are seen associated with lantern elements in BMNH
EE3735. They are relatively short and needle-like, with a
small base and no cortex. The spines are solid, not hollow.

REMARKS. The lantern of Orthopsis was described from late
Cretaceous specimens of O. sanfilippoi Checchia-Rispoli
(here treated as a synonym) by Serra (1934), who gave only a
sketchy illustration of the apparatus, but pointed out its
keeled teeth. Serra, and later Mortensen (1943: 11) described
the latern as camarodont. However, this is clearly not the
case since the epiphyses are not fused together to form a
brace for the tooth. Instead both the hemipyramid and the

A.B. SMITH

PLATE 3

Figs 1-9 Orthopsis miliaris (d’Archiac). 1-3, BMNH EE3740; 1, oral; 2, apical; 3, lateral; all x 2. Jebel Rawdah, section 3, bed 9. 4, 5, 7,
BMNH EE3725; 4, apical, x 2; 5, lateral, x 2; 7, apical disc, x 4. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m
of the Simsima Formation. 6, 8, 9, BMNH EE3733; 6, ambital ambulacrum, x 6; 8, ambital interambulacrum, x 6; 9, peristomial region,

x 6. Jebel Rawdah, section 1, bed 3.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

epiphysis extend adaxially towards the tooth as in diadema-
toid and stirodont lanterns (Jensen 1981, figs 31, 32). The
epiphyses are well-separated and although they may actually
reach the tooth and support it, they remain far apart. The
tooth is undoubtedly keeled and similar to those of stirodonts
and camarodonts.

The phylogenetic position of orthopsids can now be clari-
fied. The structure of their epiphyses and hemipyramids is
primitive for euechinoids as a whole, thus there are no
grounds for treating orthopsids as camarodonts. The keeled
teeth and solid spines place orthopsids as acroechinoids and
more derived than either diadematoids or echinothurioids.
However, the simple acrosaleniid-style of ambulacral plate
compounding and the perforate, non-crenulate nature of the
primary tubercles differentiate it from all other acroechi-
noids. They are best considered as an early plesion of the
Acroechinoidea.

Orthopsis morgani Cotteau & Gauthier (1895), from the
late Senonian of southern Iran (PI. 2, figs 4, 5) is treated as a
synonym.

Order CALYCINA Gregory, 1900
Family SALENIIDAE Agassiz, 1838
Genus SALENIA Gray, 1835

Salenia nutrix Peron & Gauthier, 1881 PI. 4, figs 1-13;
Pl. 17, figs 4, 5; Figs 15, 16A—-E, G, H

1881 Salenia nutrix Peron & Gauthier, in Cotteau, Peron &
Gauthier: 167, pl. 18, figs 4-10.

1895 Salenia cossiaea Cotteau & Gauthier: 83, pl. 13, figs
13-19.

1902 Salenia cossiaea Cotteau & Gauthier; Gauthier: 149,
pl. 18, fig. 12.

1928 Salenia cossiaea Cotteau & Gauthier; Lees: 659.

1932 Salenia lamberti Checchia-Rispoli: 6, pl. 2, figs 1-5.

1969 Salenia geometrica Agassiz; Devries: 167, pl. 1, fig. 1,
pl. 4, figs 1-3.

1989 Salenia cossiaea Cotteau & Gauthier; Ali: 398, fig. 2.6.

MATERIAL. 34 BMNH ~ EE3622-EE3654,

EE3656.

OCCURRENCE. In the western Oman Mountian sections stud-
_ ied this species was found as follows:
Jebel Aqabah; lowest few metres of the Simsima Formation
(2).
Jebel Buhays, section 1; loose in scree derived from lowest
few metres of the Simsima Formation (7).
Jebel Buhays, section 3: loose in scree (2).
_ Jebel Faiyah, section 1: bed 8 (2).
_ Jebel Rawdah, section 1; beds 3 and 4 (4); loose in scree (1).
Jebel Rawdah, section 2; beds 6-8 (10); bed 11 (2); bed 13
(2); loose in scree (3).
Jebel Rawdah section 3; bed 2b (1); bed 10 (1).
Jebel Rawdah section 4; bed 10 (1).

specimens,

This species also occurs in the Upper Campanian of Nafun,
3-5 km west of Surayr, near Dumq, Oman (Skelton ef al.
' 1990). It was originally described from the ‘Etage dordonien
_inferieur, couches a Heterolampas maresi’ (Late Campanian
or early Maastrichtian) of Algeria. Later Gauthier (1902)
described the same species from the Campanian of Tunisia.
Cotteau & Gauthier described a very similar form from the

139

Senonian of Persia (Iran) under the name Salenia cossiaea
(Pl. 4, figs 4-6). Finally, Devries (1967) described this species
from Turkey under the name Salenia geometrica Agassiz.

DIAGNOsIS. Rather inflated species of Salenia with seven to
eight interambulacral plates at test diameters of 23-28 mm.
Apical disc relatively small, circular and flat with periproct
and suranal equal in size. Ocular I insert or exsert. Numerous
small pits developed along all apical disc sutures in larger
specimens, but largest along genital/ocular plate boundaries.
Interradial granular zone well developed in specimens greater
than 23 mm test diameter.

DESCRIPTION. Tests range in diameter from 7 to 26 mm. Test
height in specimens larger than about 10 mm is 60-76% of the
diameter (mean 65%, N = 19; Fig. 15). The test is thus rather
inflated in profile and in some specimens almost subglobular
(Pl. 4, fig. 3).

The apical disc is rather flat and only rises very slightly
towards the apex. Its diameter is 42-55% of the test diameter
(mean = 46%, N = 19). It is subcircular in outline. The
suranal plate is relatively large, on average about 25% of the
apical disc diameter. It is similar in size to the genital plates.
The periproct is approximately the same size as the suranal
plate, or very slightly larger, being on average 27% of the
apical disc diameter. In some specimens ocular 1 is strongly
exsert and forms the posterior wall of the periproct, but in
other specimens ocular 1 is insert and separated from the
periproct (Figs 16A-E). Approximately half of the specimens
have the ocular plate insert. There is a slight elevation
towards the periproct edge, but no true rim is developed. All
plates are smooth and unornamented. The sutures are usually
incised and may have a series of small pits along their length
(Pl. 4, figs 1, 4, 7, 11). The ocular/genital plate boundaries
always have pits that are more prominent than the rest.
Gonopores are present on genital plates from approximately
10 mm diameter.

Ambulacra are narrow and only very slightly sinuous
towards the apex (PI. 4, fig. 13). They expand adorally to
form a short phyllode. Plating is strictly bigeminate through-
out. There are 45 pore-pairs in a column at 13 mm test
diameter, rising to 78 at 26 mm test diameter (Fig. 15). Each
compound plate has a primary tubercle that forms a contigu-
ous row adjacent to the pore-pairs (Figs 16G, H). The
perradial zone is narrow, but there is a single secondary
tubercle on each compound plate and a single zig-zag row of
miliaries also. On the oral surface up to 12 pore-pairs become
crowded into a short phyllode in the largest specimens. There
are approximately 15 or 16 ambulacral plates opposite an
ambital interambulacral plate in individuals of 20-26 mm test
diameter.

Interambulacra remain relatively broad throughout. There
are six interambulacral plates in a column from 10-16 mm test
diameter, rising to eight by about 25 mm diameter (Fig. 15).
Plates at the ambitus are slightly wider than tall and the
primary tubercle lies towards the adradial margin, leaving a
relatively broad interradial zone in specimens larger than
about 22 mm diameter. Primary tubercles are imperforate
and crenulate and are surrounded by five or six secondary
tubercles (non-contiguous) (Pl. 4, fig. 13). The interradial
zone has miliary tubercles from about 15 mm test diameter
upwards and in larger specimens this forms a broad and
distinctive band that runs almost to the peristome edge.

The peristome is on average 48% of the test diameter

140 A.B. SMITH

PLATE 4

Figs 1-13 Salenia nutrix Peron & Gauthier. 1-3, 13, BMNH EE3646; 1, apical, x 2; 2, oral, x 2; 3, lateral, x 2; 13, ambital detail, x 5.
Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 4-6, B18723a, Museum d’Histoire
Naturelle, Paris; topotype material from the Morgan Collection of Salenia cossiae Cotteau & Gauthier; 4, apical; 5, lateral; 6, oral; all x 3.
Senonian, Kala é Melek, Iran. 7-9, BMNH EE3651; 7, apical; 8, oral; 9, lateral; all x 2. Jebel Buhays, lowest 2 m of the Simsima
Formation. 10-12, BMNH EE3652; 10, lateral; 11, apical; 12, oral; all x 2. Jebel Buhays, lowest 2 m of the Simsima Formation.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

141

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Fig. 15 Biometric data for Salenia nutrix Peron & Gauthier (squares) and S. microprocta sp. nov. (crosses).

- across. It is hardly invaginated and has feeble buccal notches

(Pl. 4, figs 2, 6, 8, 12).

REMARKS. This species somewhat resembles S. geometrica
- Agassiz, but differs from that species in having a relatively
smaller apical disc and a larger periproctal opening. It also
has much less pronounced pits developed at triple suture
| junctions on the apical disc. Although there are clearly two
| morphological forms, those with ocular 1 insert and those
) with ocular 1 exsert, the apical discs are in all other respects
identical and the two forms co-occur in the various sections. I
therefore treat the two forms as variants of the same species.
ka form described by Checchia-Rispoli (1932a) as S. lam-
berti from the Maastrichtian of Libya has all the characteris-
tics of S. nutrix, except that ocular 1 is reportedly always

insert. It too is treated as part of this same species complex.
S. nutrix resembles S. loveni (Cotteau) in having a rather flat
and smooth apical disc and in having expanded phyllodes
adorally. However, it is very different both in the relative size
of the apical disc (which in S. loveni occupies most of the
upper surface) and in the coarseness of the tuberculation. S.
maxima Arnaud and S. belgica Lambert are also similar in
having fine sutural pitting but also have relatively much larger
apical discs than is seen in S. nutrix.

Fig. 16 Camera lucida drawings of plating in Salenia. A-E, apical
discs of Salenia nutrix Peron & Gauthier; A, BMNH EE3646; B,
BMNH EE3654; C, BMNH EE3647; D, BMNH EE3651; E,
BMNH EE3627. F, apical disc of Salenia microprocta sp. nov.
BMNH EE3657. G, H ambital ambulacral plating of Salenia
nutrix Peron & Gauthier; G, BMNH EE3647; H, BMNH
EE3638. I, ambital ambulacral plating of Salenia microprocta sp.
nov., BMNH EE3657. Scale bars; A-D = 5 mm; E, F = 1 mm;
G-I = 1mm.

PIS, es 1S skies 15;
16F, I

Salenia microprocta sp. nov

Type. Holotype and only known specimen, BMNH EE3657.
OCCURRENCE. Jebel Huwayyah, section 2: bed 1.

DIAGNOsIS. A small, flattish Salenia with a relatively large,
flat, pentagonal apical disc and a small oval periproct.
Ambulacral tubercles are not contiguous and at the ambitus
may be separated by granules.

DESCRIPTION. Test 11-4 mm in diameter and 5-1 mm in
height (45% of test diameter). Depressed in profile with a
broad flat base and top. The ambitus is uniformly rounded.
The apical disc is flat and pentagonal in outline, with the
angles radial. The disc is 55% of the test diameter in length.
The genital plates are approximately as broad as long and are
all similar in size (Fig. 16F). Ocular plates protrude slightly.
The suranal plate is relatively large, whereas the periproct is
small (Pl. 5, fig. 1), being only 17% of the apical disc
diameter along the plane of bilateral symmetry. There is no
lip to the periproct. Sutural pits are present at all plate triple
junctions and also mid-way between these junctions. Small
gonopores are present.

The ambulacra are slightly sinuous adapically, becoming
straight adorally and expanding towards the peristome. There
are 33 pore-pairs in a column and 16 primary tubercles. All
plates are compound and bigeminate. At the ambitus the
ambulacral width is 1-5 mm with the perradial tuberculate
zone making up 1-0 mm of this width. Primary tubercles are
relatively small and are not contiguous with their neighbours.
There is a single row of miliary tubercles perradially and, at
the ambitus, there is also a single row of miliaries between
successive primary tubercles (Pl. 5, fig. 3; Fig. 161). Small
phyllodes are developed adorally.

A.B. SMITH

Interambulacra are composed of five plates in each col- |
umn. Each has a single primary tubercle surrounded by six
widely-spaced scrobicular tubercles. There is a single row of
miliary tubercles down the interradius.

The peristome is very slightly invaginated and is 50% of the
test diameter in diameter. There are feeble buccal notches.

REMARKS. No other Salenia species has ambulacra with
primary tubercles separated by rows of granules. S. micro-
procta is also easily distinguished from small specimens of S.
nutrix by the very small size of its periproct and the relatively
large area occupied by the apical disc.

Order ARBACIOIDA Gregory, 1900
Family GONIOPYGIDAE Smith & Wright, 1993
Genus GONIOPYGUS Agassiz, 1838

Goniopygus arabicus sp. nov. Pl. 6, figs 3-10; Pl. 7, figs
1, 3,5, 6; Figs 17, 18A, B; DEG

1972 Goniopygus superbus Cotteau & Gauthier; Kier: 68,
pl. 42, figs 1-3.

1989 Goniopygus superbus Cotteau & Gauthier; Ali: 401,
in. 2 (2),

Types. The holotype is BMNH EE4012, paratypes are
BMNH EE3983-84, EE39896, EE3992, EE3997, EE4005,
EE4007, EE4015, EE4017 and EE4019.

MATERIAL STUDIED. Forty-four specimens of which the type
series was used in the biometric analysis.

OCCURRENCE. In the western Oman Mountains this species

is found at the following levels:

Jebel Buhays, section 1: loose in scree derived from the
lowest few metres of the Simsima Formation (29): bed 12
(1).

Jebel Buhays, section 2: loose in scree derived from the basal
few metres of the Simsima Formation (4).

Jebel Buhays, section 3: basal 1 m of the Simsima Formation
(4).

Jebel Thanais: lowest 1 m of the Simsima Formation (2).

Jebel Faiyah, section 1: bed 6 (1 fragment).

Jebel Rawdah, section 2: bed 4 (1): beds 6-8 (10); bed 10 (1);
bed 11 (1).

Jebel Rawdah, section 3a, bed 2 (2).

Elsewhere the species has been reported from the late
Campanian of the Rihyad district of Saudi Arabia (Kier
1972).

DIAGNOSIS. A species of Goniopygus with relatively narrow
ambulacra with a single small secondary tubercle on each
compound plate, a trigonal periproct with, predominantly,
three perianal tubercles and apical disc plating that is smooth
and unornamented. Gonopores lie on the genital plates.

DESCRIPTION. Tests range in size from 25 to 41 mm in
diameter and are circular in outline. Test height is 53-62% of
test diameter (mean = 57%, SD = 3-0%, N = 9: Fig. 17).
Both the base and top are flat in profile and the ambitus lies
at approximately mid-height (PI. 6, figs 4, 5, 7). The apical
disc plates are elevated above the corona.

The apical disc is flat, large and prominent. It occupies
35-43% of the test diameter (mean = 40%, SD = 3-3%, N=
9). Genital plates are pointed distally and the gonopore opens

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

) PLATE 5

| Figs 1-3 Salenia microprocta sp. nov. BMNH EE3657, holotype; 1, apical; 2, oral; 3, lateral; all x 6. Jebel Huwayyah, section 2, bed 1.
‘Figs 4-10, 12 Mimiosalenia quinquetuberculata gen. et sp. nov. Jebel Faiyah, section 1, bed 7. 4, 12, BMNH EE3981, holotype; 4, apical,

| 4; 12, lateral, ambulacrum detail, x 8. 5, 6, BMNH EE3982, paratype, apical; 5, x 3; 6, x 6. 7-9, BMNH EE3978, paratype; 7, lateral; 8,
apical; 9, oral; all x 3. 10, BMNH EE3980, paratype; apical, x 3.

|Fig. 11 Glyphopneustes hattaensis Ali. BMNH EE4027, detail of ambital region, lateral view, x 8; Jebel Thanais, lowest 2 m of the Simsima

| .

Formation.

144 A.B. SMITH

PLATE 6

Figs 1,2 Goniopygus superbus Cotteau & Gauthier. L12680, Lambert Collection, Geology Department, Université de Paris VI, Paris;1, |
apical; 2, lateral; both x 2. Senonian, Derré-i-Chahr, Iran.

Figs 3-10 Goniopygus arabicus sp. nov. 3, 4, 10 BMNH EE4005, paratype; 3, apical; 4, lateral; both x 2; 10, ambital detail, x 5. Jebel
Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 5, 8, BMNH EE4017, paratype; 5, lateral; 8, :
apical; both x 2. Jebel Rawdah, section 2, bed 6. 6, 7, BMNH EE4012, holotype; 6, apical, x 1-6; 7, lateral, x 2. Jebel Rawdah, section 3, |
bed 2. 9, BMNH EE3983; apical disc, variety with five perianal tubercles, x 5. Jebel Buhays, section 1; loose in the scree derived from the |

lowest 3 m of the Simsima Formation. |

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

24
22
E
E 20
—
‘3
S 18
©
fe
+ 16
©
Ee
14
12
30 40 50
Test diameter (mm)
20
@ Interambulacrum oe
gs Ambulacrum
-
e
E
~
on
3
=
2s 10
=
©
~
©
cS
~
2
=
0
Test diameter (mm)
20
@ Apical disc
@ Periproct
-_
£
E
—
-
©
z
«
fa)
Test diameter (mm)
Fig. 17 Biometric data for Goniopygus arabicus sp. nov.

beyond the tip of the apical disc platform, though still within
the genital plate (Figs 18A, B). Ocular plates are relatively
large and are insert. All plates are flat and smooth, without
Ormamentation. The periproct is oval and lies slightly poste-
rior of centre. It is approximately 9-12% of the test diameter
in width along the anterior-posterior axis. In the great
Majority of specimens the opening is trigonal and there are
three perianal tubercles on genital plates (Pl. 7, fig. 1).
However, there is a single specimen ( BMNH EE3983) that
| has five perianal tubercles (PI. 6, fig. 9) and one that has four,
thus the number of tubercles is not invariant.
| The ambulacra are 13-14% of the test diameter in width at
the ambitus. All plates are compound and trigeminate, with a
demiplate and two full elements (Fig. 18D). The two major
elements carry a single primary tubercle and the upper also

Number of primary tubercles
in a column

145

Peristome diameter (mm)

30 40 50
Test diameter (mm)

30
@ Interambulacrum

®@ Ambulacrum

N
oO

=
oO

30 40 50
Test diameter (mm)

N le)
- © fo)

Number of pore-pairs in a column
a
°

wn
fo)

30 40 50
Test diameter (mm)

has a perradial secondary tubercle in addition (Fig. 18E). The
perradial tuberculate zone thus is composed of an outer series
of primary tubercles alternating with a distinct inner series of
secondary tubercles (Pl. 7, fig. 3). All tubercles are imperfo-
rate and non-crenulate. Below the ambitus pore-pairs are
small and oval and become crowded close to the peristome to
form a relatively well-developed phyllode (PI. 7, fig. 5). At
the ambitus pore-pairs become markedly more elongate and
the individual pores more widely separated. Individual pores
in a pore-pair are distinctly conjugate in larger individuals.
There are no sphaeridial pits. There are 56 pore-pairs and 19
primary tubercles in a column at 25 mm test diameter, rising
to 77 pore-pairs and 25 primary tubercles at 41 mm test
diameter (Fig. 17).

Interambulacra are broad and each plate carries a large

146 A.B. SMITH

PLATE 7

Figs 1,3, 5,6 Goniopygus arabicus sp. nov. 1, 2, BMNH EE4012, holotype; apical disc, x 5. Jebel Rawdah, section 3, bed 2. 3, 6, BMNH
EE4015, paratype; 3, detail of ambulacrum at ambitus, x 5; 6, detail of interambulacrum at ambitus, x 3. Jebel Thanais, lowest 1 m of the
Simsima Formation. 5, BMNH EE3997, adoral detail, x 3-5. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the
Simsima Formation.

Figs 2,4 Goniopygus superbus Cotteau & Gauthier. L12680, Lambert Collection, Geology Department, Université de Paris VI, Paris; 2, |
apical disc, x 4; 4, detail of ambulacrum at ambitus, x 6. Derré-i-Chahr, Iran.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

147

Fig. 18 Camera lucida drawings of Goniopygus species. A, B apical disc plating of Goniopygus arabicus sp. nov.; A, BMNH EE4017; B,
BMNH EE4005S. C, F, apical disc plating and ambital ambulacral tuberculation of Goniopygus superbus Cotteau & Gauthier, L12680,
Geology Department, Universite de Paris VI; Senonian, Derre-i-Chahr, Iran. D, E, ambital ambulacral plating, Goniopygus arabicus sp.
nov.; D, BMNH EE4017; E, BMNH EE4005. G, G. arabicus sp. nov., adoral plating, ambulacrum to left, BMNH EE3986. Scale bars = 1

mm.

primary tubercle. At the ambitus interambulacral width is

42-46% of the test diameter. There are 8 plates in a column

at 25 mm diameter, rising to 11 at about 40 mm test diameter

(Fig. 17). Primary tubercles are stout, imperforate and non-

crenulate at the ambitus, but reduce sharply in size adapically

so that the top three or four tubercles are very small. At the
-ambitus they are surrounded by about 6 widely-spaced scro-
bicular tubercles (Pl. 7, fig. 6). Down the interradius there is
a single column of secondary tubercles on each plate.
Adorally both columns of plates reach the peristomial margin
_and there is no primordial plate (Fig. 18G).

The peristome is slightly invaginated and is 37-44% of the
test diameter in diameter (mean = 42%, SD = 2:6%, N = 7).
Buccal notches are relatively small and poorly differentiated,
with only a weak rim.

REMARKS. This species was described under the name
Goniopygus superbus Cotteau & Gauthier by Kier (1972) and
Ali (1989). It differs from that species in several important
respects. Firstly, the apical disc of G. superbus has a larger
periproctal opening and has either five, or rarely four peria-
nal tubercles (PI. 6, figs 1, 2; Pl. 7, fig. 2; Fig. 18C). These
perianal tubercles face upwards and the adjacent portions of
the genital plates are raised in the form of a very characteris-
tic stellate rim. The radial portions of this rim project
upwards as blunt denticles. Furthermore, the apical disc
plates of G. superbus are covered in fine granular ornament.
The gonopores in all specimens studied open not in the
genital plates, but within the interambulacral plates some one
or two plates distant from the apical disc (Fig. 18C). Finally,
he ambulacral tuberculation affords an easily distinguishable

character: in G. superbus the ambulacra are wide and the
inner series of secondary tubercles almost as large as the
primary tubercles, whereas in G. arabicus the ambulacra are
narrow and the secondary tubercles very much smaller (com-
pare Pl. 7, fig. 3 and Fig. 18E, with PI. 7, fig. 4 and Fig. 18F).
Finally, in G. superbus there are zones of small miliary
granules separating successive tubercles which are totally
absent in G. arabicus.

Genus MIMIOSALENIA gen. nov.
TYPE SPECIES. Mimiosalenia quinquetuberculata sp. nov.

DIAGNOsIS. A goniopygid with a perianal tubercle on each
genital plate and pits along apical disc sutures. Ambulacra
bigeminate except adorally where occasional simple plates
are intercalated.

OCCURRENCE. Known only from the late Cretaceous (Maas-
trichtian) of Jebel Faiyah, section 1, western Oman Moun-
tains.

REMARKS. This genus is closely related to Goniopygus on
account of its distinctive apical disc plating. The five gonop-
ores lie immediately beyond the genital plates and open in the
interambulacra. The genital plates of the apical disc have
perianal tubercles identical to those of Goniopygus and the
stellate ridge surrounding them is very reminiscent of that
seen in G. superbus Cotteau & Gauthier. However, there are
well-developed sutural pits both at triple junctions and mid-
length along the plate sutures on all apical disc plates, which

148
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E
E
cd
i=
2
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is
~
n
©
=
6 8 10 12 14
Test diameter (mm)
2.5
C=
E
E
= 72{0)
©
~
©
:
= 1.5
~
0
°
i=
a
= 10
©
a
0.5
3 4 5 6 7 8
Apical disc diameter (mm)
8 @
B Peristome ) ‘
@ Apical disc i.
E
E
o
-
o
=
fs
(S)
6 8 10 Uz 14

Test diameter (mm)

Fig. 19 Biometric data for Mimiosalenia quinquetuberculata sp. nov.

are often seen in saleniids but are never seen in Goniopygus.
Another major difference between Goniopygus and Mimios-
alenia is that the ambulacra of Goniopygus are trigeminate or
occasionally quadrigeminate, whereas those of Mimiosalenia
are strictly bigeminate, except close to the peristome where
plating tends to become unigeminate. Again, bigeminate
plating is typical of saleniids. All ambulacral plates in Mimi-
osalenia reach the perradius and are approximately equal in
size, whereas in Goniopygus there is a demiplate and the
other two plates in each triad are unequal in size.

Despite the similarities to Salenia, Mimiosalenia lacks a
suranal plate and has the highly characteristic apical disc
structure of a goniopygid. For this reason I believe it to be a
derived goniopygid that has developed sutural pitting and

Width at ambitus (mm)

Number in column

A.B. SMITH

® Ambulacrum
6 @ Interambulacrum oe

6 8 10 12 14 16
Test diameter (mm)
40
B® Pore-pairs a i
@ Primary ambulacral tubercles
2" :

10
Test diameter (mm)

12 14 16

bigeminate plating through the loss of the demiplate in each|
triad.

Mimiosalenia quiquetuberculata sp. nov P1. 5, figs 4-10,
12; Figs 19, 20)7

Types. Holotype, BMNH EE3981; paratypes, BMNH|
EE3971, EE3974, EE3977-78, EE3980, EE3982,
EES014-17.

MATERIAL STUDIED. Biometric data is taken from the type
series. In addition there are six other specimens.

OCCURRENCE. The species is known only from bed 7, section
1b, Jebel Faiyah, western Oman Mountains.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

DIAGNOsIS. As for generic diagnosis: apical disc with five
perianal tubercles. Gonopores lie just outside the genital
plates in the interambulacral areas.

DESCRIPTION. Tests range from 7-2 to 15-5 mm in diameter
and are circular in outline. Test height is 42-57% of test
diameter with the flatter tests possibly representing slightly
crushed specimens (Fig. 19). In profile the apical disc is very
slightly conical and the sides uniformly rounded so that the
ambitus lies at about mid-height.

The apical disc is relatively large and raised above the
coronal plates (PI. 5, fig. 7). It is 48-57% of the test diameter
across (Fig. 19) with the periproct more or less centrally
positioned. Ocular plates are relatively large and square-
ended. The five genital plates border the periproct and are
bluntly pointed distally (Figs 20A, B). All plates are smooth
and unornamented. Each genital plate has a single perianal
tubercle adjacent to the periproct. The genital plates are
raised to form a stellate rim surrounding these perianal
tubercles (PI. 5, figs 4, 6; Figs 20A, B). All five genital plates
are approximately the same size. Gonopores open beyond
the genital plates and are found immediately adjacent in the
interambulacral zones (Fig. 20B). They are only present in
the larger specimens, ca. 14 mm diameter. The periproct is
subcircular in outline and measures 9-15% of the test diam-
eter in width (Fig. 19).

Ambulacra are relatively broad and slightly concave perra-
dially. They are 11-17% of the test diameter in width at the
ambitus. From the apex to below the ambitus plating is
strictly bigeminate (Pl. 5, fig. 12; Fig. 20C), with each
element reaching the perradius (Fig. 20D). Each pair of
elements bears a large primary tubercle. Close to the peris-
tome there are occasional simple elements interspersed, each
with a large primary tubercle. There are 31-32 pore-pairs and
‘16-17 primary tubercles in a column at 11-5-13 mm test
‘diameter, rising to 37 pore-pairs and 19 primary tubercles at
‘15-5 mm test diameter (Fig. 19). The perradial zone of
tuberculation is very broad and contains a mixture of second-
ary tubercles and granules, two or three abreast (PI. 5, fig. 12;
Fig. 20C). Adorally the pore-pairs are slightly more widely
separated and there is no phyllode development whatsoever.
Ambulacra hardly taper either adorally or adapically. There
\are no sphaeridial pits.

Interambulacra are 38-44% of the test diameter in width at
the ambitus. There are seven plates in a column at 6-2 mm
‘est diameter, rising to eight or nine at 15 mm test diameter.
Bach plate carries a large primary tubercle which is non-
vrenulate and imperforate. The most adapical two are gener-
illy very much smaller than the remainder. The primary
ubercles have six scrobicular tubercles, three on each side,
hat are more or less contiguous (PI. 5, fig. 12). The interra-
lial zone is broad and slightly concave. It is occupied by a

eries of miliary tubercles, two to each plate (four abreast).
‘here is no primordial plate adorally and both columns reach
€ peristomial border.

The peristome is circular, slightly invaginated and occupies
640% of the test diameter. Buccal notches are small and
idistinct (PI. 5, fig. 9).

Spines, lantern and perignathic girdle all unknown.

|
|EMARKS. The biserial nature of the ambulacra and the
laracteristic apical disc structure make this species easy to

stinguish from any other described here.

149

Fig. 20 Camera lucida drawings of plating in Mimiosalenia
quinquetuberculata sp. nov. A, apical disc, BMNH EE3981; B,
apical disc, BMNH EE3982; C, ambulacrum, from apical disc
(top) to peristomial margin (bottom), BMNH EE3981; D, ambital
ambulacral plating, BMNH EES5014~-17. Scale bars = 1 mm.

Family GLYPHOPNEUSTIDAE Smith & Wright, 1993
Genus GLYPHOPNEUSTES Pomel, 1883

Glyphopneustes hattaensis Ali, 1992 PI. 5, fig. 11; Pl. 8,
figs 1-12; Figs 21, 22

1992a Glyphopneustes hattaensis Ali: 68, fig. 3.

Types. Holotype, the figured specimen, housed in the Geo-
logical Museum, University of Al Ain, United Arab Emir-
ates.

MATERIAL STUDIED. 82 specimens, of which biometric data
was taken from the following 30: BMNH EE3909, EE3913,
EE3915, EE3919-20, EE3923-24, EE3926, EE3930-31,
EE3934, EE3936-38, EE3940, EE3942-43, EE3945-47,
EE3949, EE3953-5S4, EE3958, EE3960, EE3967-70.

OCCURRENCE. The type material all comes from Jebel Raw-

dah. This species was collected at the following levels:

Jebel Huwayyah, section 2: bed 1 (1).

Jebel Faiyah, section 1: bed 2 (1); bed 5 at base (1); bed 8 (9);
loose approximately 10 m above base of the Simsima
Formation (1).

150

Jebel Buhays section 1; loose, derived from lowest few
metres of the Simsima Formation (50).

Jebel Thanais: lowest 2 m of the Simsima Formation (6).

Jebel Rawdah, section 1; bed 4 (1).

Jebel Rawdah, section 2; beds 9/10 (2); bed 11 (5); bed 13 (1);
loose (6).

Jebel Rawdah, section 3; bed 2 (2); bed 5 (1).

Jebel Rawdah, section 4; bed 4 (1).

DESCRIPTION. Tests range in diameter from 10 to 30 mm,
with the great majority around 18-24 mm diameter (Fig. 21).
The test is circular in outline and depressed in profile, with a
rounded ambitus, although some specimens are slightly more
conical. Test height is 39-57% of the diameter (mean = 50%,

—_
wn

Test height (mm)
°

30
Test diameter (mm)

10 20

swt
8 - os

Peristome diameter (mm)

10 20 30
Test diameter (mm)
14
@ Interambulacrum AY
s Ambulacrum

12 ee

€ -*
yy

10
£ «3
a °
ome o%,
3
E
«
~
i=
~
1
=

Test diameter (mm)

Fig. 21 Biometric data for Glyphopneustes hattaensis Ali.

A.B. SMITH

SD = 4:0%, N = 30; Fig. 21). The ambitus lies at about |
mid-height.

The apical disc is highly sculpted and occupies 25-42% of
the disc diameter (mean = 32%, SD = 1-2%, N = 30). It is
proportionally larger in small individuals (Fig. 21). The
periproct is large and central, occupying 30-50% of the apical
disc diameter (mean = 40%, SD = 4-9%, N = 28). It is oval |
in outline with smoothly rounded edges (PI. 8, figs 1, 2). The
apical disc is dicyclic and all five genital plates are approxi- |
mately equal-sized (Fig. 22B). There is a rim surrounding the
periproct which bears a large central tubercle and two lateral
tubercles. The gonopores open at the outer edge of the
genital plates. The madreporite has a horse-shoe-shaped zone
of madrepores that open around its margin (Fig. 22B). |

25
gs Ambulacrum (compound)
E @ Interambulacrum
=
°
1S)
@
=
n
©
~
w
a
Sa
i)
he
4
E
=]
z

(2)
lo}

wn
jo)

+
°o

Number of pore-pairs in a column

30
10 20 30 40
Test diameter (mm)
10
w Apical disc
@ Periproct r
8 a
a = =
€ & os z
E S f*,*
a a
© r
-_
ae oe
s Se
a 9 Ad ONE Ie
e e
2F eo y
°

10 20

30 40

Test diameter (mm)

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

LATE 8

jigs 1-12 Glyphopneustes hattaensis Ali. 1, BMNH EE3930; apical disc, x 6. Jebel Buhays, section 1; loose in the scree derived from the
lowest 3 m of the Simsima Formation. 2, 8, BMNH EE3943; 2, apical disc, x 5; 8, apical, x 2. Jebel Buhays, section 1; loose in the scree

| derived from the lowest 3 m of the Simsima Formation. 3, 6, 9, BMNH EE3915; 3, ambital detail of ambulacrum, x 6. 6, ambital detail of

_interambulacrum, 4; 9, peristomial detail, x 6. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima

Formation. 7, BMNH EE3958; apical, x 2. Jebel Faiyah, section 1, 2 m above the base of the Simsima Formation. 10, 11, BMNH EE4027;

}

10, oral, x 2; 11, lateral, x 2. Jebel Thanais, lowest 2 m of the Simsima Formation. 4, 5, 12, BMNH EE3945; 4, apical; 5, oral; 12, lateral;
all x 3. Jebel Buhays, section 1, loose in the scree.

152

Ocular plates are heart-shaped. The sutures are all deeply
incised and there are typically three small tubercles along the
ocular/genital plate boundary within these depressed regions.

Ambulacra are 13-17% of the test diameter in width at the
ambitus (Fig. 21). All plates are trigeminate and there is a
single demiplate in each triad (Figs 22A, C). The two full
elements bear a single large primary tubercle that is imperfo-
rate and non-crenulate. The upper element is smaller than
the lower. Towards the peristome the demiplate has a shallow
pit immediately perradial to the pore-pair, which marks the
site of a sphaeridium (PI. 8, fig. 3; Fig. 22D). There are four
or five of these in each column. Pore-pairs are uniserially
arranged and not noticeably enlarged ambitally and adapi-
cally. There is no pore crowding whatsoever close to the
peristome. There are around 33 pore pairs and 11 primary
tubercles in a column at 11 mm test diameter, rising to about
60 pore-pairs and 21 primary tubercles (Fig. 21). There is a
single row of scattered miliary tubercles down the perradius.

Interambulacra are 39-46% of the test diameter in width at
the ambitus (mean = 44%, SD = 1-8%, N = 30). There are 9
plates in a column at 10 mm test diameter, rising to 12 at 30
mm test diameter (Fig. 21). Ambital plates are much wider
than tall and each bears a single large primary tubercle that is
non-crenulate and imperforate (Pl. 8, fig. 6). The most
adapical two or three plates have significantly smaller
tubercles than the rest. The primary tubercles have three
scrobicular tubercles on either side, but have confluent
areoles within each column. Down the interradius there are
two or three irregular rows of scattered secondary and miliary
tubercles, forming a relatively broad granular zone (PI. 8, fig.
6). Adorally both columns of plates reach the peristome and
there is no primordial plate.

The peristome is rather small and not at all invaginated. It
is 33-43% of the test diameter across (mean = 38%, SD =
2:9%, N = 27), proportionally smaller in larger individuals
(Fig. 21). Buccal notches are relatively shallow.

REMARKS. Ali (1992a) gave a detailed description of this
species but based on only six specimens. The large number of
well-preserved specimens now to hand allows a detailed
biometric description of this species for the first time. Ali
specifically stated that sphaeridial pits were lacking in this
species, yet in well-preserved specimens such pits can be
seen. The difference between this species and the Cenoma-
nian G. problematicus rest almost entirely on the apical disc
ornamentation and the absence of sutural pits on the interam-
bulacral plates of G. hattaensis. Although Fell & Pawson
(1966) placed Glyphopneustes in the family Hyposaleniidae,
and were followed by Smith & Wright (1990), it is now
evident from the material available that Glyphopneustes is an
arbacioid, and has been placed in its own family Glyphop-
neustidae by Smith & Wright (1993). It has the characteristic
perianal tubercles and apical disc structure of that family, and
also has a similar style of ambulacral plate compounding. The
sphaeridial pits are shallow and clearly convergent with those
in Hyposalenia.

A.B. SMITH

Fig. 22 Camera lucida drawings of plating in Glyphopneustes
hattaensis Ali. A, Ambulacral plating, from ocular plate (top) to
peristome margin (bottom), BMNH EE3926; B, apical disc,
BMNH EE3912; C, ambital ambulacral plating, BMNH EE3915;
D, adoral ambulacral plating, peristomial border at bottom,
BMNH EE3915. Scale bars = 1 mm.

Family ARBACIIDAE Gray, 1835
Genus CODIOPSIS Agassiz, 1840

Codiopsis lehmannae sp. nov PI. 9, figs 1-2; Pl. 12, figs
1-3; Fig. 23

DERIVATION OF NAME. After Mrs C. Lehmann, the finder of
the holotype.

Types. Holotype, BMNH EE5033; paratypes (both incom-
plete test fragments), BMNH EE3439, EE3440.

OCCURRENCE. One specimen comes from Bed 15, section 1,
Jebel Buhays, a second comes from bed 10 (top), section 2,
Jebel Rawdah. The third specimen was found loose in the
basal scree at Jebel Rawdah, section 2, and is almost certainly
derived from the lowest few metres of the Simsima Formation
(beds 3-10).

DESCRIPTION. The holotype is 18-3 mm in diameter and 8-2
mm in height (45% of the diameter). The two other speci-
mens are larger, but incomplete, and by estimation would
have been around 35—40 mm in test diameter. The base 1s flat,
the upper surface domal, and the ambitus is very sharp and at
the base. The apical disc is preserved in BMNH EE3440 and
EE5033. It is dicyclic and firmly fixed to the corona (PI. 12,
fig. 1; Fig. 23C). The periproct is oval and 1-7 mm in
diameter in the 18-3 mm diameter individual (9-3%).

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

Fig. 23 Camera lucida drawings of plating in Codiopsis lehamannae
sp. nov. A, ambulacral plating (one column only) from
supra-ambital region (top) to peristomial margin (bottom),
BMNH E3439; B, aboral ambulacral plating, a little above the
ambitus, BMNH EE34339; C, apical disc (incomplete) showing
parts of two genital plates and two ocular plates, BMNH EE3440.
Scale bars = 1 mm.

Ambulacra are narrow and parallel-sided above the ambi-
'tus, where they are trigeminate. Plate compounding is
arbaciid in style with two demiplates (Fig. 23B). There is a
single calcitic pustule on each triad (PI. 9, figs 1, 2). Pores are
small and widely-spaced, possibly conjugate (though preser-
vation is too poor to be certain). Below the ambitus the
| pore-pairs reduce in width markedly (Fig. 23A) and the
-ambulacra widen into extensive phyllodes. The pore-pairs
adorally are crowded and circular in outline with well-
_ developed periporal muscle attachment areas. Compounding
becomes polygeminate in a complex fashion (Fig. 23A) and
there is a single large tubercle on each compound plate.
Interambulacral plates are geniculate at the ambitus and
) adoral portions bear a single primary tubercle. This arrange-
/ment creates a row of slightly downward-facing tubercles
) along the ambitus (PI. 9, figs 1, 2). Adradial portions of these
/plates have fine secondary tuberculation. Adradial to the
| primary tubercles and continuing adapically along the adra-
)dial margin are very large calcite pustules. These reduce in
‘size considerably above the ambitus and appear to continue
adapically at least most of the way to the apical disc. The
|remainder of the adapical plates may have a pitted epistroma
\(traces are seen in BMNH EE34339), but the test is not well
preserved.

‘REMARKS. There is no doubt as to the generic placement of
‘these specimens, on account of their distinctive pustules and
acid style ambulacral compounding. Their sharp ambitus,
flat base, extensive phyllodes and ambital ring of interambu-
lacral tubercles are distinctive and are features unknown in
other species. It differs from Codiopsis brunei Lambert, from

the Maastrichtian of Maastricht, in being very much larger

153

and in lacking well-developed aboral pustules. It comes closer
to C. disculus Peron & Gauthier (and its synonyms C.
stephensoni Cooke and C. fontei Vidal) from the late
Campanian/Maastrichtian of Algeria, southern Spain, Sene-
gal, Brazil and southern U.S.A., but differs from that species
in having a more depressed profile, sharper ambitus and
more distinct ambital ring of large tubercles. In C. lehmannae
the primary interambulacral tubercles form a peripheral row,
whereas in C. disculus the tubercles form a V-shaped
arrangement extending adorally. Finally, in C. Jehmannae the
peristome appears highly scalloped.

Genus HATTOPSIS Ali, 1992

Hattopsis sphericus Ali, 1992 Pl. 9, figs 3-8; Pl. 10, figs
1, 2, 4; Fig. 24, 25B-D, 26B, 27A

1992b Hattopsis sphericus Ali: 694, fig. 3.

Types. Holotype 910401 in the Museum of the Geology
Department, University of Al Ain, United Arab Emirates.
Paratype 910402.

MATERIAL STUDIED. 73 specimens, of which the biometric
data was taken from the following: BMNH EE3658, EE3663,
EE3690, EE3692-95, EE3698, EE3702, EE3705,
EE3707-20.

OCCURRENCE. The species was described from Jebel Raw-

dah (?section 1) by Ali (1992). Material collected in situ is as

follows:

Jebel Faiyah, section 1: bed 7 (1); bed 8 (pycnodont level)
(8).

Jebel Rawdah, section 1: bed 3 (22); bed 4, mostly towards
top and base of bed 5 (44).

Jebel Rawda, section 2: bed 11 (2); loose in scree at base of
section (1).

DIaGNosis. A spherical arbacioid with a reticulate epistro-
mal ornamentation. Every third ambulacral pore-pair
reduced in size. Two interambulacral tubercles on each plate.
Peristomial rim elevated as a lip interradially.

DESCRIPTION. Tests range from 12-0 to 20-8 mm in diameter
(Fig. 24) and are circular to rounded pentagonal in outline.
Test height is 60-83% of the diameter (mean = 74%, SD =
6:5%, N = 16), and tests are globular in profile with a small
base and apex (PI. 10, fig. 2).

The apical disc is dicyclic and occupies 23-33% of the test
diameter (mean = 27%, SD = 2:4%, N = 14). The periproct
is oval in outline and 9-12% of the test diameter in diameter.
Ocular plates project slightly beyond the ring of genital plates
(Pl. 10, fig. 2; Figs 25B, C). Genital plates are large and flat
except around the periproctal margin where they are raised to
form a rim. All genital plates are similar in size. The
madreporite pores extend over most of genital plate 2.
Well-preserved specimens show a reticulate pattern of ridges
and pits (Pl. 10, fig. 2).

Ambulacra are 20-24% of the test diameter in width at the
ambitus (Fig. 24). They are compound throughout with
trigeminate plating. Close to the peristome both upper and
lower elements are demiplates, but elsewhere it is only the
upper element that is a demiplate (Fig. 26B). This demiplate
has a pore-pair that is very much smaller than those on the
other two elements. Each triad has a single large primary

A.B. SMITH

PLATE 9
Figs 1,2 Codiopsis lehmannae sp. nov. BMNH EE34339, paratype; 1, oral; 2, lateral; both x 3. Jebel Buhays, section 1, bed 15.

Figs 3-8 Hattopsis sphericus Ali. 3-5, BMNH EE3710; 3, apical; 4, oral; 5, lateral; all x 3. Jebel Rawdah, section 2, bed 11. 6-8, BMNH
EE3712; 6, apical; 7, oral; 8, lateral; all x 3. Jebel Rawdah, section 2, bed 11.

Figs 9-11 Hattopsis paucituberculatus sp. nov. BANH EE3683, holotype; 9, apical; 10, oral; 11, lateral; all x 3. Jebel Buhays, section 1;
loose in the scree derived from the lowest 3 m of the Simsima Formation.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 155

PLATE 10

)Figs1,2,4 Hattopsis sphericus Ali. BANH EE3660; 1, adoral detail showing sphaeridial pits at perradius and peristomial lip, 6; 2, apical
disc, X 6; 4, detail of ambital interambulacrum, x 6. Jebel Faiyah, section 1, bed 11.

\Fig.5 Noetlingaster paucituberculatus (Noetling). BMNH EE3680, juvenile, apical view, x 4 (see also PI. 11, Figs 4, 5). Jebel Buhays,
section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation.

‘Figs 3,6 Hattopsis paucituberculatus sp. nov. 3, BMNH EE3672, paratype, adapical detail, x 7. Jebel Faiyah, section 1, bed 2. 6, BMNH

_ EE3682, paratype; ambital detail, interambulacrum to left, x 6. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of

| the Simsima Formation.

156

tubercle. There is a small secondary tubercle on the perradial
margin of the lower element but other miliaries and second-
ary tubercles are absent. The perradius is ornamented with a
reticulate pattern of ridges and shallow pits (PI. 10, fig. 4).
Close to the peristome there are up to four sphaeridial pits
arranged uniserially down the perradius (PI. 10, fig. 1). There
are 33 pore-pairs and 10 primary tubercles in an ambulacral
column at 12 mm test diameter, rising to 45-50 pore-pairs and
14-16 primary tubercles at 20-21 mm diameter (Fig. 24). No
phyllodes nor any hint of pore-pair crushing is seen towards
the peristome, and the pore-pairs themselves become much
smaller adorally (Pl. 9, figs 4, 7; Fig. 26B).

Interambulacra are 36-40% of the test diameter in width at
the ambitus. Plates are wider than tall and carry two small
primary tubercles placed towards the adradial margin (PI. 10,
fig. 4; Fig. 27A). The tubercles in each pair are contiguous

V8 rs H. sphericus

@ H. paucituberculatus

=
a
a

+
a

@
B C4
_ a
=» 4
~~, wo % s

Test height (mm)
o oN

10 12 14 16 18 20
Test diameter (mm)
60r gH sphericus

E @ H. paucituberculatus

6 50

8.40

-—

° 30

=20

10 12 14 16 18 20
Test diameter (mm)
16 8 H. sphericus

@ H. paucituberculatus

Number of Interambulacral plates

16 18
Test diameter (mm)

Fig. 24

A.B. SMITH

but are well separated from pairs of tubercles on other plates.
There are no secondary or miliary tubercles, the remainder of
the plate being covered in the same reticulate ornament of
ridges and pits. At the peristome edge the interambulacra are
thickened to form a distinct lip (Pl. 10, fig. 1). There is a
single T-shaped primordial interambulacral plate forming the
border to the peristome (Fig. 25D). There are 11 interambu-
lacral plates in a column at 12 mm diameter, rising to 14 or 15
at 20-21 mm test diamater (Fig. 24).

The peristome is not at all invaginated and is 38-48% of the
test diameter across (mean = 43%, SD = 3-1%, N = 13).
Buccal notches are very slight and the raised interambulacral
rim forms the most prominent feature (PI. 10, fig. 1).

REMARKS. Ali (1992b) gave a detailed description of this

species but new features reported here for the first time |

g H. sphericus
@ H. paucituberculatus

Apical disc diameter (mm)

16 18
Test diameter (mm)

12 14 20 22

9p eH. sphericus
@ H. paucituberculatus

12 14 16 18 22

Test diameter (mm)

Biometric data for Hattopsis sphericus Ali and Hattopsis paucituberculatus sp. nov.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

include the perradial uniserial row of sphaeridial pits, the
large T-shaped primordial interambulacral plate and the
extensive development of reticulate ornamentation. The pri-
mordial plate, the umiserial and perradially positioned
sphaeridial pits and the style of ambulacral compounding
clearly place Hattopsis in the Arbaciidae. The reduction of
one pore-pair in each triad and the presence of primary
interambulacral tubercles above the ambitus separate Hattop-
sis from Codiopsis, which it resembles. Hattopsis comes most
close in structure and appearance to juveniles of Noetlin-
gaster, but differs from them in having fewer interambulacral
tubercles at comparable sizes, and in being more globular in
profile.

Hattopsis paucituberculatus sp. nov. Pl. 9, figs 9-11; PI.
10, figs 3, 6; Pl. 11, fig. 9; Figs 24, 25A, 26A, 27B

Types. Holotype EE3683, paratypes, EE3682, EE3678,
EE3688, EE3684-85.

MATERIAL STUDIED. There are 22 specimens in addition to
_ the types. Biometric data is based on the following speci-
_ mens: BMNH EE3672, EE3675, EE3678-79, EE3682-85,
_ _EE3688.

‘ ties and horizons:

_ Jebel Faiyah, section 1: bed 2 (12).

| Jebel Buhays section 1: in scree derived from lowest beds of
the Simsima Formation (11); base of bed 12 (1).

Jebel Buhays, section 2: loose in scree, derived from lowest 3
m of the Simsima Formation (1).

| Jebel Buhays, section 3: basal beds of the Simsima Formation

(2).

| Jebel Aqabah: bed 1 (1).

| Jebel Rawdah, section 2, bed 8 (1); loose in scree derived

from lowest part of section (1).

_ DIAGNOsIs. A species of Hattopsis with only a single inter-
| ambulacral tubercle on each interambulacral plate at all sizes.

DESCRIPTION. Tests are 6-2 to 18-2 mm in diameter and
circular to rounded pentagonal in outline (Pl. 9, figs 9-11).
| Test height is 58-83% of test diameter (mean = 71%, SD =
-6-7%, N = 9) and in profile the test appears globular.
The apical disc is dicyclic, like that of H. sphericus (Fig.
25A). It is 28-37% (mean = 31%) of test diameter in
diameter and there is an elevated rim around the periproct
margin (Pl. 10, fig. 3). Gonopores are generally rather larger
and more oval than those of H. sphericus.
. Ambulacra are 20-25% of the test diameter in width at the
-ambitus and the ambulacral compounding is as in H. spheri-
cus (Fig. 26A). There are 27 pore-pairs and 8 tubercles in a
column at 10-7 mm test diameter, rising to 44 pore-pairs and
14 tubercles at 18:2 mm test diameter (Fig. 24). Aborally
from the ambitus, the pore-pair on the upper demiplate in
each triad is greatly reduced in size (Pl. 10, fig. 6; Fig. 26A).
Adorally all pore-pairs become small and they remain unise-
\rial to the peristome edge.
| Interambulacra are 36-39% of the test diameter in width.
Each plate carries a small primary tubercle, situated towards
\the adradial margin. (PI. 10, fig. 6; Fig. 27B). The remainder
‘of the plate is covered in fine reticulate ridges and pits. There
‘are no secondary tubercles developed, even in the largest
specimens.

The peristome is 43-47% of the test diameter in diameter

Fig. 25 Camera lucida drawings of plating in Hattopsis. A-C, apical
discs; A, H. paucituberculatus sp. nov, BMNH EE3672; B, H.
sphericus Ali, BMNH EE3692; C, H. sphericus Ali, BMNH
EE3681. D, H. sphericus, BMNH EE3693, adoral interambulacral
plating, peristomial edge at base. Scale bar = 1 mm.

Fig. 26 Camera lucida drawings of ambulacral plating in Hattopsis.
A, H. paucituberculatus sp. nov., complete ambulacrum from
apical disc (top) to peristomial margin (bottom); BMNH EE3678;
B, H. sphericus Ali, ambulacrum from close to apical disc (top) to
peristomial margin (bottom); BMNH EE3659. Scale bars = 1
mm.

and has the usual interambulacral lip and shallow buccal
notches.

REMARKS. This species resembles H. sphericus in all details
except that it has only a single interambulacral tubercle on
each plate, rather than the two found on all specimens of H.
sphericus greater than 12 mm. Although I have included

A.B. SMITH

PLATE 11

Figs 1-3 Noetlingaster emiratescus Ali. BMNH EE3285; 1, oral; 2, apical; 3, lateral; all x 1. Jebel Rawdah, section 4, bed 12.

Figs 4-7 Noetlingaster paucituberculatus (Noetling). 4, 5, BMNH EE3680 (juvenile); 4, lateral; 5, oral; both x 4. Jebel Buhays, section 1;
loose in the scree derived from the lowest 3 m of the Simsima Formation. 6, BMNH EE3286 (juvenile), lateral view, x 4. Jebel Rawdah,
section 2, bed 11. 7, BMNH EE3282, detail of ambital region, adoral towards top and interambulacrum to left, x 4.

Fig.8 ?Noetlingaster sp. BMNH EE36839, lateral, x 5. Jebel Huwayyah, section 2, beds 2-7.

Fig. 9 Hattopsis paucttuberculatus sp. nov. BMNH EE3682, lateral view, x 3. Jebel Buhays, section 1; loose in the scree derived from the

lowest 3 m of the Simsima Formation.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

some specimens smaller than 12 mm in this species, they
could possibly turn out to be juveniles of H. sphericus.
However, the majority of specimens are larger than 12 mm
and the distribution of the two species is also not the same.
Whereas H. sphericus is found predominantly at Jebel Raw-
dah, section 1, beds 34 and also in beds higher up the
succession (bed 11) in section 2, as well as at the pycnodont
level (bed 8) at Jebel Faiyah, H. paucituberculatus consis-
tently occurs further down in the succession (basal beds of
Jebel Buhays and Jebel Faiyah, and bed 8 in Jebel Rawdah,
section 2). The two species are thus stratigraphically discrete.

Genus NOETLINGASTER Vredenburg, 1911

TYPE SPECIES. Protechinus paucituberculatus Noetling, 1897
by original designation.

REMARKS. Noetlingaster has previously been classified in the
Stomechinidae on account of its imperforate tuberculation
and dicyclic apical disc (e.g. Fell & Pawson 1966). However,
it has a single large primordial interambulacral plate at the
adoral end of each interambulacrum and a tuberculation style
very similar to that of Hattopsis. Primordial interambulacral
plates this well-developed are known only in the Arbaciidae.
Finally, juvenile forms of Noetlingaster are extremely similar
in appearance to Hattopsis, differing only in having a more
depressed test profile and more interambulacral tubercles at
corresponding sizes. Given the marked similarity between
Hattopsis and Noetlingaster in apical disc, tuberculation,
interradial epistroma and pore-pair development, there seem
strong grounds for placing Noetlingaster as an arbaciid.
There are nine named species of Noetlingaster, all from the
late Cretaceous. The type species N. paucituberculatus (Noet-

Fig. 27 Camera lucida drawings of ambital plating in Hattopsis and
Noetlingaster? sp.: interambulacral plate on the left, ambulacral
plate on the right. A, Hattopsis sphericus Ali, BMNH EE3693; B,
Hattopsis paucituberculatus sp. nov., BMNH EE3683; C,
Noetlingaster? sp., BMNH EE3689.

159

ling, 1897) comes from horizon 4 (?Maastrichtian) in the Des
Valley, Mari Hills, West Pakistan. Other species are: N.
emiratescus Ali (1989), described from Jebel Rawdah, west-
ern Oman mountains; N. sanfilippoi Checchia-Rispoli (1930),
N. millosevichi Checchia-Rispoli (1930) and N. lamberti
Checchia-Rispoli (1930), all from the Maastrichtian of Gebel
Misid, Tripolitania, Lybia; N. globulus Devries (1967) and N.
hemisphericus Devries (1967), both from the Maastrichtian of
Kahta, Turkey; N. monteili Gauthier (1901) from the ‘Senon-
ian’ south of Bilma, Algeria; and N. boulei Lambert (1906)
from the Maastrichtian of Marohite, Madagascar. Devries
(1967) reviewed previous species and discussed their diagno-
sis. He laid particular emphasis on the arrangement of
interambulacral tubercles, recognizing two groups: those with
a single row of interambulacral tubercles on each plate, and
those with a double row. He pointed out that the actual
number of tubercles in each row varied ontogenetically and
also according to position on the test.

All these species are closely related because of their highly
distinctive ambulacral plate compounding and pore-pair
arrangement (see below). However, all have been erected on
the basis of very few specimens, often simply the holotype.
Thus the intraspecific variability has never been properly
documented. In addition, the relatively thin test means that
specimens are rarely well preserved. The large collection of
specimens from Jebel Rawdah allows ontogenetic variability
to be assessed in this species complex for the first time.

There are significant differences in size between the species
that have been erected, and these may account for some of
the morphological variation described. N. boulei is described
from the smallest specimen, only 49 mm in diameter. N.
globulus, N. hemisphericus and N. emiratescus are all based
on specimens ranging from 65-83 mm. The remainder are
described from large individuals between 95 and 120 mm in
diameter (Fig. 28). As many of the characters previously used
for species discrimination can be shown to vary with size in
the Omani populations, it is important that similar-sized
individuals are compared when differentiating species.

Shape differences were used by Devries (1967) to separate
some species. He identified species as either ‘subconical’ or
‘rounded convex’. However, there seems to be considerable
variation in this feature within the sample described here, and
thus the character has little worth. It is also difficult to use in
practice since specimens are often crushed or distorted during
preservation.

Ambulacral differences have also been used for diagnosing
species. N. boulei for example has just one primary tubercle
on ambulacral plates, and this is found on alternate plates
only. However, as Lambert’s (1906) photographs show,
ambulacral tuberculation is not consistent. Adoral compound
plates all have a single primary tubercle, whereas, adapically,
tuberculation is more irregular with occasional plates lacking
primary tubercles. Considering the small size of N. boulei one
would not expect more than a single tubercle to be devel-
oped, by comparison with the Omani sample. Furthermore,
tuberculation is very irregular in the Omani population,
especially adapically from the ambitus where it is relatively
common to find primary tubercles missing from ocassional
plates. Such irregular ambulacral tuberculation characterises
all species and from about 60 mm test diameter upwards all
species have plates bearing two or three tubercles irregularly
arranged. Note that although Gauthier (1901) describes N.
monteili as having eight rows of ambulacral tubercles, his
figures show only four irregular columns and the ambulacral

160

Number of primary
tubercles in row

10
2 [a
G
oe
—-
5 ee se
2
see
e Wee 2 B
ao
@

A.B. SMITH

e N paucituberculatus
0 N emiratescus

OH =double row of
tubercles present

20 25 30

Plate width (mm)

Fig. 28 Biometric data from Noetlingaster species. Type specimens for species are plotted as follows: B = N. boulei Lambert; E = N.
emiratescus Ali (two syntypes); G = N. globulus Devries; H = N. hemisphaericus Devries; L = N. lamberti Checchia-Rispoli; Mi = N.

millosevichi Checchia-Rispoli; Mo = N. monteili Gauthier.

structure does not differ from that seen in equivalent-sized
Omani specimens.

Most emphasis has been placed on interambulacral tuber-
culation for differentiating between species. Devries (1967)
recognized two species groups: those with just a single row of
tubercles to each interambulacral plate and those with a
double row. He also distinguished between those with a
naked interradial zone of fine granulation and those with
primary tubercles extending more or less to the interradius.
Unfortunately, the Omani specimens also show considerable
variation in these features. The majority simply have a single
row of primary tubercles extending more or less to the
perradius. From the ambitus adapically there may or may not
be a naked zone free of primary tubercles and when present
this zone may or may not extend below the ambitus. In many
specimens it is difficult to observe tuberculation interradially
because of weathering, and the same is probably true for the
specimens on which other species are based. Single rows of
tubercles are characteristic of N. paucituberculatus, N. mon-
teili, N. boulei and N. globulus, whereas N. sanfilippoi, N.
millosevichi, N. hemisphaericus and N. emiratescus all have
two or three additional tubercles forming a second row
adradially, and N. lamberti has a full double row of tubercles
developed on all plates. All tuberculation styles, except that
seen in N. lamberti, are encountered in the Omani sample.
As can be seen from Fig. 28, there is a strong correlation
between the number of tubercles on a plate and the plate
width (which is proportional to test diameter). Furthermore,
it is primarily the larger specimens that have a second row of
tubercles developed adradially. However, it is not completely
size-dependent since the 69 mm diameter specimen EE3279
has a well developed secondary row of tubercles whereas
others of that size do not.

In conclusion, the variation observed within the 20 reason-
ably well-preserved specimens from the western Oman
mountains is almost as great as that observed between the
nine described species, each based on one or a few specimens

only. The presence or absence of an interradial naked zone
and the extent of this zone when present, the number of
interambulacral tubercles in a row in proportion to the plate
width and the development of a second row of primary
tubercles to a plate are all variable. Similar variation of
tuberculation has been found in both Turkey (Devries 1967)
and Libya (Checchia-Rispoli 1930), but was used to distin-
guish ‘species’.

By contrast the analysis of variation amongst Omani speci-
mens suggests that tuberculation pattern may not be so rigidly
developed. It does, however, support some subdivision of the
genus. When tuberculation style is plotted against test diam-
eter, two distinct growth trajectories emerge for Omani
specimens (Fig. 28). In some specimens there are numerous
densely-spaced tubercles with secondary tubercle rows
present, even at 60 mm test diameter. Other specimens have
more widely-spaced primary tubercles and only develop a
secondary row of adradial tubercles on ambital plates at a
very large size. Although these may simply represent end
members of a continuous spectrum, the small sample does
not indicate any significant overlap. Those with many
tubercles also always lack a granular interradial zone,
whereas those with relatively fewer tubercles typically have a
naked zone. However, the development of this naked zone is
quite variable, with some specimens showing a broad zone
extending almost to the peristome, and others with the zone
restricted to a narrow wedge-shaped area adapically. Mea-
surements taken from the types of all nine species are plotted
for comparison. The type species N. paucituberculatus falls
into the growth series of forms with fewer tubercles, as do the
types of N. millosevichi, N. boulei, N. monteili, N. sanfilip-
poi, and N. lamberti. The more densely tuberculate form
corresponds to N. emiratescus and may also include N.
globulus and N. hemisphaericus, although both plot in an
intermediate position. (This may be due to inaccuracies in the
magnifications for the camera lucida drawings of interambu-
lacral plates given approximately by Devries (1967, pl. 2)).

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

Plotting those with a secondary row of tubercles to inter-
ambulacral plates shows that the secondary row appears at
about 15 mm plate width in N. globulus/N. emiratescus,
whereas it only develops at around 22 mm plate width in the
N. paucituberculatus group.

The one anomolous specimen is BMNH EE3282, which
has numerous small interambulacral tubercles, but a wide
interradial granular zone. It falls in between the two growth
series in Fig. 28.

In conclusion, only two species of Noetlingaster are recog-

nized here:

(1) Forms to ca. 125 mm diameter with granular interradial
zone developed adapically or throughout and a single
row of relatively widely spaced interambulacral
tubercles. Adradially, a second shorter row of tubercles
is developed in larger specimens. These forms occur at
Jebel Rawdah, section 2 from bed 11 to 14 and corre-
spond to the form first described as N. paucituberculatus
(Noetling).

(2) Forms to ca. 125 mm with a second row of interambulac-
ral tubercles present adradially from about 60 mm test
diameter, and no naked zone. At comparable sizes there
are more interambulacral tubercles in a row than in the
first species. This form occurs at Jebel Rawdah section
1, but at higher levels in the section than N. paucituber-
culatus. It has been described from the Oman Moun-
tains under the name N. emiratescus Ali.

_Noetlingaster paucituberculatus (Noetling, 1897) Pl. 10,
fig. 5; Pl. 11, figs 4-7; Figs 28, 29, 30A—D, F, H, 31

11897 Protechinus paucituberculatus Noetling, 1897: 16, pl.

2, fig. 3, pl. 3, fig.1.

11898 Noetlingia paucituberculata Noetling; Lambert: 126.

1901 Noetlingia Monteili Gauthier: 191, pl. 3, figs 1-3.

/ 1906 Noetlingia Boulei Lambert: 11, pl. 2, fig. 7.

(1911 Noetlingaster paucituberculata Noetling; Vredenburg:

46.

Noetlingaster Millosevichi Checchia-Rispoli: 14, pl. 2,

figs 1, 2, pl. 4, figs 3, 4.

1930 Noetlingaster Sanfilippoi Checchia-Rispoli: 6, pl. 1,

| fig. 1, pl. 3, fig. 2, pl. 4, fig. 2.

1930 Noetlingaster Lamberti Checchia-Rispoli: 20, pl. 1, fig.
2, pl. 3, fig. 1, pl. 4, fig. 1.

Types. The type is the 95 mm diameter specimen described
by Noetling (1897) from the late Cretaceous of the Mari Hills,
West Pakistan.

MATERIAL STUDIED. 19 relatively complete tests and 2 test
fragments, including juveniles. The following ten specimens
were used in the biometric analysis: BMNH EE3267, 3269,
3271-75, 3280, 3282 and 3286.

‘OCCURRENCE. In the western Oman mountain area N. pau-

‘cituberculatus was found at the following levels:

Jebel Rawdah section 1, base of bed 4 (juvenile).

Jebel Rawdah, section 2: bed 11 (3, plus fragments); bed 14

(5, plus fragments); bed 19 (2); bed 25 (4); bed 26 (2).

Jebel Buhays, section 1: loose in scree derived from lowest

| beds of the Simsima Formation (3, including one complete
13 mm diameter juvenile).

Outside the eastern Arabian peninsula this species is

161

recorded from western Pakistan, Algeria, Libya and Mada-
gascar.

DESCRIPTION. Tests range in diameter from 15 to 126 mm.
They are more or less circular in outline, but very slightly
depressed both interradially and adradially. Test height is
54-79% of test diameter (mean = 66%, SD = 7:2%, N = 10)
and juveniles tend to have more depressed tests than adults.
In profile the test is subconical, with a broad base, narrow
apex and low ambitus (PI. 11, figs 4, 6). The ambitus lies at
about one quarter of the test height above the base.

The apical disc is relatively small, occupying only 13-16%
of the test diameter in medium to large individuals (mean =
14%, SD = 1:2%, N = 6). It is proportionally larger in small
individuals, occupying 19% of the test diameter in the 15 mm
diameter specimen (Fig. 29). Disc plating is dicyclic (Figs
30A, B). The madreporite is very much larger than other
genital plates and is densely covered in small pores. Each
genital plate has a large oval gonopore which may be sur-
rounded by a slight rim. The ocular plates are small and
pentagonal, each with a small ocular pore. The periproctal
opening is large and oval.

Ambulacra are straight and taper adapically. At the ambi-
tus their width is 16-20% of the test diameter. Ambulacral
plates are trigeminate throughout with a highly distinctive
style of. compounding (Figs 30C, D, F, H). The lowest
element is large and occupies the full width. There is a
smaller demi-plate above, which always carries a large pore-
pair adradially and an even smaller, fully occluded element
above that (Figs 30D, F). This arrangement is found along
the entire length of the ambulacrum in all medium to large
individuals, except at the very apex, where the occluded plate
may reach the adradial suture. In smaller individuals both of
the smaller elements are demiplates (Fig. 30H), while in
BMNH EE3286, a juvenile of only 15 mm test diameter (Fig.
30F), the upper element extends to the perradial suture.
There are no phyllodes nor any pore crushing towards the
peristome. Each element carries a pore-pair, but only on the
large primary element are these well-developed throughout
(Pl. 11, fig. 7). The pore-pairs on the small occluded upper
element are always reduced to rudimentary structures and in
places may simply be represented by a single pore. The
middle element usually has well-developed pore-pairs, but in
larger individuals around the ambitus these may also be very
much reduced in size (Fig. 30F). Pore-pair differentiation is
hardly developed in the 15 mm diameter juvenile. Each
compound plate usually carries a single primary tubercle
close to the pore-zone. However, tuberculation is irregular
and occasionally two tubercles occur to a plate. In the largest
individuals most ambulacral plates carry two adradial primary
tubercles, slightly offset on alternate plates. The perradial
band is devoid of large tubercles above the ambitus, but this
zone is relatively narrow, typically only some 15% of the
interambulacral width. There are approximately 112 pore-
pairs in a column at a test diameter of 32 mm, rising to around
180 at 80 mm test diameter (Fig. 29).

Interambulacral width at the ambitus is 41-47% of the test
diameter. Plates are wide and low, and are slightly taller than
ambulacral plates. There are 36 interambulacral plates in a
column at 82 mm test diameter. The primary tubercle at the
centre of the plate is set on a slight keel which runs down the
midline of each column and is particularly prominent adapi-
cally. There are multiple small tubercles on each plate, three
in the smallest individual (PI. 11, fig. 4), rising to 11 in the

A.B. SMITH

162

PLATE 12 |

Figs 1-3 Codiopsis lehmannae sp. nov. BMNH EES033, holotype; 1, apical; 2, oral; 3, lateral; all x 4. Jebel Buhays, section 1; loose in the
scree derived from the lowest 3 m of the Simsima Formation.

Figs 4-7 Actinophyma spectabile Cotteau & Gauthier. BMNH EE3601; 4, apical, x 3; 5, oral, x 3; 6, lateral, x 3; 7, detail of ambital
plating, x 6. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

+ a [es]
fo) jo) (>)

Test height (mm)

N
jo)

0) 20 40 60 80 100 120 140

Test diameter (mm)

140

Number of pores in column
a

30 40 50 60 70 80 90

Test diameter (mm)

0 10 20 30
Interambulacral plate width at ambitus (mm)

@ Ambulacrum A
@ Interambulacrum

BSS
fo}

Width at ambitus (mm)
Ww
io)

20
10
(0)
10) 20 40 60 80 100 120 140
Test diameter (mm)
30
@ Peristome
® Apical disc
=
E
it
®
-~
o
E
&
6
0 20 40 60 80 100 120

Test diameter (mm)

Fig. 29 Biometric data for Noetlingaster paucituberculatus (Noetling) and N. emiratescus Ali.

_ largest. These are arranged in a single, slightly arcuate row
(Pl. 11, fig. 7), but the larger individuals may have two or
three additional tubercles forming an upper row near the
adradial suture. All tubercles are imperforate and non-
crenulate and have relatively small mamelons and extensive
areoles. Sutures between all plates are finely denticulate.

The peristome is very slightly sunken and is strongly

_ indented by wide buccal notches. Peristome diameter is

20-31% of test diameter (mean = 27%, SD = 4:0%, N = 7).

REMARKS. This species is distinguished from N. emiratescus
_ by its interradial granular zone, which may be small and
| deveioped only adapically, or may be broad and extend
_ adorally. Within a single population from bed 14, section 2,
| Jebel Rawdah, the development of this granular zone was

highly variable, with some having only a narrow adapical
wedge of granules, others having a broad band extending
adorally. It also differs from N. emiratescus in having fewer
interambulacral tubercles at comparable sizes (Fig. 28).
Smaller individuals (diameters less than ca. 100 mm) have
single rows of tubercles on interambulacral plates, but a
second row is present in larger specimens.

The species was first described by Noetling (1897) on the
basis of a single specimen from West Pakistan. Gauthier
(1901) later described a smaller, incomplete specimen of this
species under the name N. monteili from the eastern Sahara
region of Algeria. Other species that are probably best
treated as synonymous are the three species described by
Checchia—Rispoli (1930) from the Maastrichtian of Libya.

164

Fig. 30 Camera lucida drawings of Noetlingaster. A-D, F, H, N.
paucituberculatus (Noetling): A, B, apical disc plating: A, BMNH
EE3269; B, BMNH EE3275; C, adapical plating in a juvenile,
BMNH EE5049; D, Ambital plating, BMNH EE3266; F, ambital
plating, BMNH EE3286; H, adapical plating, BMNH EE3283. E,
G, N. emiratescus Ali, BMNH EE3285; E, adapical plating; G,
ambital plating. Scale bars = 1 mm.

Fig. 31 Camera lucida drawings of plating in a juvenile
Noetlingaster paucituberculatus (Noetling), BMNH EE3680. A,
Apical disc; B, ambital plating, ambulacrum to the left,
interambulcaral plate on right; C, adoral interambulacral plating,
showing the single primordial interambulacral plate. Scale bar = 1
mm.

There is a general increase in size up the section at Jebel
Rawdah, with small to medium-sized individuals found in the

A.B. SMITH

lower beds and only large individuals towards the top in the
deeper water facies.

Noetlingaster emiratescus Ali, 1989 P\. 11, figs 1-3; Figs
28, 29; 30E, G

1989 Noetlingaster emiratescus Ali: 398, Fig. 2 (3-5)

Types. The types of N. emiratescus are three specimens and
six fragments in the collections of the Geology Department,
United Arab Emirates University, Al Ain.

MATERIAL STUDIED. Three specimens, BMNH EE3279,
EE3284-85.

OCCURRENCE. All specimens, including the type material,
come from Jebel Rawdah. The specimens reported here were
collected from bed 4, Jebel Rawdah section 1 (1) and bed 13,
Jebel Rawdah section 4 (2).

DIAGNOsIS. A Noetlingaster with smaller and denser inter-
ambulacral tubercles than other species, and completely
lacking a naked interradial zone.

DESCRIPTION. Tests range from 68-5 to about 105 mm in
diameter. In outline the interradii and adradii are slightly
depressed. Test height is 58-71% of test diameter and in
profile the test is inflated and subglobular. The ambitus lies
about one third of test height above the base.

The apical disc occupies 11% of the test diameter and has
the same arrangement of plates as N. paucituberculatus.
Ambulacral structure is also more or less identical, with the
pore-pair on the occluded plate rudimentary almost to the
apex (Figs 30E, G). The middle pore-pair is also rudimentary
from the ambitus adorally. Ambulacral tubercles are irregu-
larly arranged and occupy the full width of the ambulacra,
one or two to a compound plate.

Interambulacral plates are slightly V-shaped, with a single
well-developed row of tubercles along the lower edge and a
second, irregular row of occasional tubercles above. There
are 19 tubercles in a row at a test diameter of 71 mm, rising to
13 at 105 mm test diameter (Fig. 28). Tubercles occupy the
full width of plates throughout and there is no granular
interradial zone as is found in N. paucituberculatus.

The peristome is small and somewhat invaginated, occupy-
ing 26% of the test diameter.

REMARKS. Ali (1989) described this species on the basis of
the material collected from Jebel Rawdah. Its small, numer-
ous interambulacral tubercles and lack of any interradial zone
of granulation serve to separate it from the co-occurring
species, N. paucituberculatus.

Noetlingaster? sp. (juvenile?) Pl. 11, fig. 8; Fig. 27C
MATERIAL. One specimen, BMNH EE3689.

OCCURRENCE. From the Loftusia beds (beds 2-5), Jebel
Huwayyah, section 2.

DIAGNOsIS. A species of arbaciid with a single primary
interambulacral tubercle on ambital and adoral interambulac-
ral plates, but smooth adapically. Ambulacral tubercles
developed to apex. Broad ambital and adapical interradial
zone ornamented with vertical riblets.

DESCRIPTION. The test is rounded pentagonal in outline and
13-2 mm in diameter. It has a broad flat base and domed

————— Eee
TTS EE — ee
a

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

upper surface. Test height is 57% of test diameter and the
ambitus lies at around 40% of test height.

The apical disc is 3-8 mm in diameter (29% of test
diameter) and is dicyclic. It is not well preserved, but the
genital plates are raised as a lip around the periproct margin
and the periproct itself is 11% of the test diameter and oval in
outline. Ocular plates protrude beyond the genital plates
slightly.

Ambulacra are 20% of the test diameter in width at the
ambitus. All plates are compound and trigeminate, but
details of plate compounding are not clear. There is clearly a
single element and a double element in each triad, but it is
impossible to tell whether one of the double elements is a
demiplate, as I suspect. All pore-pairs are equally well-
developed. There is no phyllode development. There are 33

* 10
©
Ee

(0)

30 40 50
Test diameter (mm)
20
8 Ambulacrum ’
@ Interambulacrum

~~
E
E
—
n
3
=

£10
@
~
a
<=
-
3
=

(0)

30 40

Test diameter (mm)

Number of Interamb. plates in column

40
Test diameter (mm)

Fig. 32 Biometric data for Phymosoma hexoaporum Lambertt.

165

pore-pairs and 10 primary tubercles in a column.

Interambulacra are 39% of the test diameter in width at the
ambitus. Each plate has a large adradial tubercle that extends
adapically to within three or so plates of the apex. Adorally
these tubercles lie subcentrally, but towards the apex they
become positioned more and more closely towards the adra-
dial suture. On ambital and adapical plates there is also a very
much smaller mamelonate secondary tubercle lying at the
centre of each plate within the zone of riblet ornamentation.
The entire interradial zone from the ambitus upwards, has a
well-developed ornamentation of vertically orientated riblets.

The peristome is 5-2 mm in diameter (40% of the test
diameter) and is not invaginated. There are no raised interra-
dial lips to the peristome margin. Buccal notches are
extremely feeble.

-_
+
o

® Total
¢@ Number arranged biserially

Number of pore-pairs in column

30 40

Test diameter (mm)

60 50 60

® Peristome

@ Apical disc
18 a

=
(>)

Diameter (mm)

=
N

=
fo)

50
Test diameter (mm)

60 40 60

166

REMARKS. This species is easily distinguished from Hattopsis
sphericus on the basis of its interambulacral ornamentation of
vertical riblets, and by its single row of primary interambulac-
ral tubercles. It also lacks the reduced diameter pore-pair in
each triad that is so characteristic of H. sphericus and H.
paucituberculatus. It differs from H. paucituberculatus in its
lower profile, less enlarged adapical pore-pairs and in its
riblet ornamentation as opposed to the pitted ornamentation
seen in H. paucituberculatus and H. sphericus. The vertical
riblet ornamentation of Noetlingaster? sp. is characteristic of
Codiopsis species such as C. doma, but Noetlingaster? sp.
differs from Codiopsis in having no sharp reduction in the size
of tubercles at the ambitus. It could possibly be the juvenile
of a very much larger Codiopsis species, such as C. stephen-
soni Cooke, but it has open gonopores, which suggests that it
is a genuinely small species. It differs from species such as
Codiopsis bruni Lambert & Thiéry (Maastrichtian of the
Netherlands) and C. disculus Peron & Gauthier (late Campa-
nian of Algeria, early Maastrichtian of Spain) in having a
smooth, pustule-free upper surface. This specimen most
likely represents a small Noetlingaster, one in which only the
primary interambulacral tubercles have formed. However,
neither of the named species of Noetlingaster shows the
distinct vertical ribbing that characterises this specimen.

Order PHYMOSOMATOIDA Mortensen, 1904
Family PHYMOSOMATIDAE Pomel, 1883
Genus PHYMOSOMA Haime, in d’Archiac & Haime, 1853

Pl. 13, figs
1-3; Figs 32, 33

1908 Phymosoma (Cyphosoma) Archiaci Cott. var: Cot-
fieeaNs il, fol, 3), ws, Il

1927 Phymosoma hexoaporum Lambert: 35, pl. 2, figs
25-27.

1933 Phymosoma Paronai Checchia-Rispohi: 11, pl. 2, figs
4-7.

Phymosoma hexoaporum Lambert, 1927

MATERIAL. Nineteen specimens of which the biometric data
was drawn from the following: BMNH EE3607-08,
EE3610-12, EE3614, EE3616, EE3618, EE3941.

OCCURRENCE. This species is found almost exclusively in the
lowest beds of the Simsima Formation at Jebel Buhays,
section 1 (16), and the immediately adjacent Jebel Thanais
(3). In addition one specimen was found loose in the scree in
Jebel Rawdah, section 3, and another in Bed 2 at Jebel
Rawdah, section 4. Cottreau (1908) recorded an identical
specimen from the late Cretaceous (?Maastrichtian) of Maro-
hita, Eastern Madagascar.

DIAGNOsIs. A Phymosoma with a single large primary inter-
ambulacral tubercle on all plates, and one or two small
adradial tubercles. Plates are composed of six or seven
elements at the ambitus and pore-pairs are biserial from
about the ambitus adapically. Interradial zones of small
granules are very well developed.

DESCRIPTION. Tests range from 23 to 50 mm in diameter and
are circular in outline. The test is depressed in profile (Pl. 13,
fig. 3), with a height that is 34-46% of the test diameter
(mean = 42%, SD = 4-4%, N = 8: Fig. 32). The ambitus lies
slightly below mid-height.

A.B. SMITH

The apical disc is caducous and is missing from all speci-
mens. The outline of the apical disc is pentagonal with angles
pointing interradially and projecting slightly further into the
posterior interambulacrum (PI. 13, fig. 2).

Ambulacra taper slightly both adapically and adorally.
They measure 22-26% of the test diameter in width at the
ambitus. All plates are polygeminate (Fig. 33). Above the
ambitus pore-pairs are biserial and plates are composed of six
to eight elements. Plate compounding is in the phymosomatid
style. At the ambitus the pore-pairs are in arcs of six or seven
in specimens more than 25 mm diameter, while subambitally
they are quinquegeminate. Immediately adjacent to the
peristome there are a couple of quadrigeminate plates. There
are no sphaeridial pits adorally. Each compound plate carries
a single large imperforate, crenulate tubercle, as large as the
adjacent interambulacral primary tubercles. This occupies
most of the plate. However, perradially there is a narrow
band of small secondary and miliary tubercles (Pl. 13, fig. 3).
Primary tubercles more or less reach the apex. There are
57-58 pore-pairs in a column at 23-24 mm test diameter,
rising to 138 at 50°mm test diameter. Biserial pores appear
immediately above the ambitus in most specimens and com-
prise 50-54% of the total number of pore-pairs in a column.
In the very largest specimen, biserial pore-pairs extend to the
subambital region.

Interambulacra are 32-38% of the test diameter in width at
the ambitus. There are 10 plates in a column at 23 mm test
diameter, rising to 18 at 50 mm test diameter (Fig. 32). Each |
plate carries a single large primary tubercle, centrally posi-
tioned. However, areoles are not contiguous, but are sepa- |
rated by a narrow band of miliary granules (PI. 13, fig. 3). |
Tubercles are imperforate and crenulate and decrease in size
gradually both adapically and adorally. There are one or two
small secondary tubercles to each plate situated close to the | _
adradial suture. In a few specimens these tubercles enlarge |
above the ambitus to about half the size of the primary |
tubercles, so as to form a secondary row. However, this is |
inconsistently developed. The interradius is broad and cov-
ered in scattered small secondary and miliary tubercles (Pl. |
13, fig. 3). In some specimens (e.g. BMNH EE3614) this |

tuberculate band is relatively dense, whereas in other speci- |

mens (e.g. BMNH EE3618) tuberculation is more scattered. | ~
Both columns of plates reach the peristomial border. ii
The peristome is very slightly invaginated and occupies |
40-47% of the test diameter. Buccal notches are small, but |
clearly incised (PI. 13, fig. 1).
Lantern elements are seen scattered inside the test in| —
BMNH EE3941. Hemipyramids are largely concealed by
sediment, but one keeled tooth is seen in cross-section. |

REMARKS. The variation encountered in the secondary}
tuberculation of this species is more marked than expected |
and is matched by a difference in the degree of tubercle] —
crenulation. In the specimens with the denser miliary tuber-|—
culation interradially, there are usually well-developed sec-
ondary tubercles along the adradial margin in the region)
immediately above the ambitus, and primary tubercles are}
only weakly crenulate. In the more usual variety, the interra-}
dial zone has more scattered tuberculation, there is no
enlarged secondary tuberculation and the primary tubercles
have a well-developed and broad crenulate platform. These
two forms may eventually prove to represent distinct species,
but as they are both found at the same locality and the
secondary tuberculation differences are not entirely consis-

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 167

PLATE 13
Figs 1-3 Phymosoma hexoaporum Lambert. BMNH EE3614; 1, oral; 2, apical; 3, lateral; all x 3. Jebel Buhays, section 1; loose in the scree
derived from the lowest 3 m of the Simsima Formation.

Figs 4,5 Actinophyma spectabile Cotteau & Gauthier. B18731, Morgan Collection, Museum d’Histoire Naturelle, Paris; 4, lateral; 5, apical;
both x 1-5. Senonian of Iran, no locality details.

|
|

168

Fig. 33 Camera lucida drawing of ambulacral plating, from apex
(top) to peristomial margin (bottom), in Phymosoma hexoaporum
Lambert, BMNH EE3617. Scale bar = 1 mm.

Fig. 34 Camera lucida drawings of plating in Actinophyma
spectabile Cotteau & Gauthier. A, adapical ambulacral plate from
a large individual, BMNH EE3599; B, supra-ambital ambulacral
plating in a juvenile, BMNH EE3601; C, adoral ambulacral
plating in a juvenile, BMNH EE3601. Scale bar = 1mm.

tent, the two forms are simply treated as varieties of the same
species here.

The species described here appears identical to the speci-
men described from the Maastrichtian of Madagascar by

A.B. SMITH

Cottreau (1908) under the name Phymosoma archiaci Cot-
teau var. It differs from true P. archiaci in having a broader,
more granular interradial zone, and in having less well- |
developed secondary tubercles on interambulacral plates. |
The species P. hexoaporum was established by Lambert i
(1927) for specimens from the Maastrichtian of Sopeira, —
province of Aragon, Spain. P. hexoaporum differed from all |
other species of Phymosoma described previously in having |
compound plates composed of six elements at the ambitus |
rather than five. Later, Checchia-Rispoli (1933) described the
same species from the Maastrichtian of Libya under the name |
P. paronai Checchia-Rispoli. This too has six or seven l)
elements in ambital ambulacral compound plates.

Genus ACTINOPHYMA Cotteau & Gauthier, 1895

Actinophyma spectabile Cotteau & Gauthier, 1895 PI.
12, figs 4-7; Pl. 13, figs 4, 5; Fig. 34)

1895 Actinophyma spectabile Cotteau & Gauthier: 98, pl.
25, figs 6-10.

1895 Cyphosoma persicum Cotteau & Gauthier: 93, pl. 25, |
figs 3-4.

1902 Actinophyma spectabile Cotteau & Gauthier;
Gauthier: 151, pl. 20, figs 7-10.

1935 Actinophyma_ spectabile Cotteau & Gauthier; ~

Mortensen: 489, fig. 287. |

Types. The holotype is the specimen described by Cotteau &

Gauthier from the upper Senonian of Endjir-kouh, Aftab
district, Iran.

MATERIAL STUDIED. Six specimens, BMNH EE3598-3603,| —
three of which are test fragments only. The following descrip-| -
tion is based on the two more or less complete specimens} ©
BMNH EE3601 (a juvenile) and BMNH EE3603 (an adult).| ~
Topotype material of A. spectabile in the Natural History| ©
Museum and in the Morgan Collection, Museum d’Histoire
Naturelle, Paris, has been studied for comparison. |

OCCURRENCE. Five of the specimens come from scree collec-
tions at Jebel Buhays, section 1 and are derived from the
lowest few metres of the Simsima Formation. Two other
specimens come from Jebel Rawdah section 3b, one from bed
8, the other from bed 9.

The species range is ‘Upper Senonian’ of southern Iran and) ~
Maastrichtian of the Oman Mountains.

DIAGNOsIS. A species of Actinophyma with strong radiating) ©
ridges on the boss of primary tubercles. Ambulacra polyploid
adapically with up to 18 pore-pairs to an ambulacral plate
arranged irregularly.

DESCRIPTION. Tests range in diameter from 21-2 mm to
approximately 90 mm and are circular in outline. Test height
is 33 mm in the larger specimen and 9-4 mm in the smalle
(ca. 35-45% test diameter). In profile the ambitus is more 0
less at mid-height and is smoothly rounded below and slightly
more conical above.

The apical disc is small; 5-8 mm diameter in the 2] m
individual (21% of test diameter) and probably only about 15
mm in the larger individual (16% of test diameter). Apical) _
disc plates are lost from all specimens and were evidently not
securely sutured to the corona. The opening is pentagonal in
outline (Pl. 12, fig. 4; Pl. 13, fig. 5).

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

' Ambulacra are almost as broad as the interambulacra,
' being 25-3 mm wide at the ambitus in BMNH EE3603, as
_ compared to the interambulacral width of 28-7 mm. Pore-
| pairs are in arcs of six on the oral surface, becoming biserial
- subambitally in the larger specimen and continuing so to the
apex (Fig. 34A). Adapical plates have up to 18 pore-pairs,
and show phymosomatid-style compounding (Fig. 34A). In
the small individual adapical pore-pairs are only just begining
to become biserial and all plates have just six or seven
_ elements (PI. 12, fig. 7; Fig. 34B). There are 10 or so plates in
» acolumn at 21 mm diameter and 18 or 19 at 90 mm diameter.
‘Each plate carries a single large primary tubercle that occu-
_ pies most of the plate. However, in the larger individual there
is a narrow adradial band of heterogeneous small secondary
and miliary tuberculation. Ambulacral tubercles are not
' contiguous, but are separated by a band of small secondary
tubercles. Areoles are strongly sculptured by radial grooves
' that extend from the base towards the platform of the boss.

; The adoral plating in the small individual (BMNH EE3601)
is noteworthy, since it comprises simple plating arranged in
triads (Fig. 34C). However, standard phymosomatoid-style
+ compounding develops subambitally, and this appears to be
simply a juvenile feature.

There are 11 interambulacral plates in a column at 21 mm
| test diameter, rising to about 18 at 90 mm test diameter. Each
) plate carries a large primary tubercle with a small imperforate
‘mamelon and a crenulate platform. Areoles at the ambitus
‘and adorally are confluent and are oval in outline. The areole
) bears strong radial grooves which extend up the sides of the
‘boss (PI. 12, figs 5, 6). These are most pronounced in the
smaller individual. Tubercles decrease in size above the
‘ambitus. The primary tubercles are centrally positioned on
\the plate and there are relatively broad adradial and interra-
\dial bands of secondary tuberculation. On plates around the
jambitus there is an enlarged secondary tubercle to the
adradial side of the primary tubercle, but otherwise second-
ary tuberculation is small and heterogeneous. The interradial
‘zone is depressed and tubercle-free towards the apex.

The peristome is relatively small and invaginated. In the 21
‘mm individual it is 7-1 mm in diameter (33% of test diam-
eter), and is proportionally smaller in the larger individual.
Buccal notches are very shallow (Pl. 12, fig. 5).

‘REMARKS. When first erected (Cotteau & Gauthier, 1895),
‘this species was based on a small individual 29 mm in
diameter, which is virtually identical to BMNH EE3601 in
morphology. In the same publication Cotteau & Gauthier
(1895: 91) erected another species, Cyphosoma persicum on
the basis of a larger, but fragmentary specimen from Derre-i-
Chahr. Subsequently, with the collection of more material,
Gauthier (1902) recognized that A. spectabile and C. persi-
‘\cum were simply different growth stages of the same species
and synonymized the two, selecting A. spectabile as the valid
ame.

The characteristic radial striation, which is so strongly
evident in juveniles, makes this an easily recognizable spe-
cies. Only one other comparable species has been described,
Actinophyma cf. A. spectabile Kier (1972: 68), from the
‘Campanian of Saudi Arabia. This differs from the Iranian
and Omani species in having deep pits developed at the
2orners of interambulacral plates on interradial sutures.

169

Test height (mm)
a N [o-)

10 20 30
Test diameter (mm)

® Ambulacrum
@ Interambulacrum e

Width at ambitus (mm)

Test diameter (mm)

@ Peristome s
10 ®@ Apical disc
E
=
©
~
©
E
ol
(=)
20 30
Test diameter (mm)
Fig. 35 Biometric data for Plistophyma asiaticum Cotteau &
Gauthier.

Genus PLISTOPHYMA Peron & Gauthier, in Cotteau,
Peron & Gauthier, 1881.

Plistophyma asiaticum Cotteau & Gauthier, 1895PI. 14,

figs 1-7; Figs 35, 36

1895 Plistophyma asiaticum Cotteau & Gauthier: 105, pl.
16, figs 11-14.

Types. The holotype is the single specimen described and
figured by Cotteau & Gauthier. It is not in the Morgan
collection in the Museum d’Histoire Naturelle, Paris.

MATERIAL STUDIED. Six specimens, BMNH EE3572-76,
EE4932. Only BMNH EE3573, which is incomplete, was

170
omitted from the biometric analysis given below.

OCCURRENCE. In the western Oman Mountains this species

was found at the following levels and localities:

Jebel Buhays, section 1: loose in scree, derived from lowest
few metres of the Simsima Formation (4).

Jebel Thanais: From the lowest metre of Simsima Formation
(2).

Jebel Rawdah, section 2, bed 4 (1).

The species was described from the Senonian of Derre-i-
Chahr, southern Iran and there are no other records.

DIAGNOsIS. A Plistophyma in which the ambital interambu-
lacral plates are very much smaller and narrower than either
adoral or adapical plates.

DESCRIPTION. Tests range in diameter from 12 to 25-5 mm
and are rounded pentagonal in outline with the angles
interradial. Tests are depressed and rounded in profile, with
the ambitus at or very slightly below mid-height (PI. 14, figs
3, 4). Test height is 33-38% of test diameter (mean = 36%).

The apical disc is large and pentagonal in outline with the
angles interradial (Pl. 14, fig. 2). It is 47-50% of the test
diameter in diameter and is not invaginated. No specimen
retains any apical disc plates but, to judge from the size and
shape of the opening, disc plating was presumably monocy-
clic.

Ambulacra are 13-16% of the test diameter in width at the
ambitus. They are narrowest ambitally and expand slightly
adapically (Fig. 36A). Plates both at the ambitus and adorally
are trigeminate with all three elements reaching the perradial
suture (Fig. 36C). The central element is the largest. Each
plate carries a single imperforate, non-crenulate tubercle
which straddles all three elements. Adorally, pore-pairs
become crowded so as to form small phyllodes (Pl. 14, fig. 6).
Immediately above the ambitus pore-pairs become biserially
arranged and primary tubercles diminish in size and do not
reach the apex (PI. 14, fig. 5). There are 48 pore-pairs in a
column (of which the most adapical 18-20 are biserially
arranged). There are 12 or 13 primary tubercles. Secondary
tubercles lie immediately adjacent to each pore-pair and may
also occur along the perradius on occasional elements.

Interambulacra are 39-44% of the test diameter in width at
the ambitus. Each plate is very wide and short and bears a
row of small, equal-sized tubercles. These are imperforate
and appear non-crenulate, but on close inspection of well-
preserved material (e.g. BMNH EE3573) there are faint
traces of crenulation to larger tubercles. At the ambitus the
interambulacral plates become very much narrower and the
size of the tubercles more or less halves (PI. 14, figs 3, 4; Fig.
36B). All plates are arranged to form a pronounced
downward-pointing V. Both columns of plates reach the
peristomial border. The peristome is 42-52% of the test
diameter in diameter and is hardly invaginated. There are
very feeble buccal notches. The perignathic girdle consists of
small auricles which do not meet above the perradius.

REMARKS. The species is distinguished from the type species
P. africanum Peron & Gauthier (Cotteau et al. 1881) by its
somewhat sharper decrease in interambulacral plate size at
the ambitus. However, the two species are very similar
indeed in other features and the Algerian and Omani-Iranian
species may eventually turn out to be conspecific. The
residual crenulation on well-preserved tubercles and the large

A.B. SMITH

Fig. 36 Camera lucida drawings of plating in Plistophyma

asiaticum Cotteau & Gauthier, BMNH EE3575. A, ambulacrum,
from apex (top) to peristomial margin (bottom); B,
interambulacrum from apex (top) to peristomial margin (bottom);
C, detail af ambital ambulacral plating. Scale bars = 1 mm.

Fig. 37 Camera lucida drawings of plating in Circopeltis? emiratus

sp. nov. A, apical disc, BMNH EE3584; B ambital ambulacral
plating, BMNH EE3582; C, ambital plating, ambulacrum to right,
interambulacral plate to left, BMNH EE3584. Scale bar = 1 mm.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

PLATE 14

Figs 1-7 Plistophyma asiaticum Cotteau & Gauthier. 1-3, 6, BMNH EE4932; 1, oral, x 5; 2, apical, x 5; 3, lateral, x 5; 6, detail of
peristomial region, x 8. Jebel Thanais, lowest 1 m of the Simsima Formation. 4, 7, BMNH EE3575; 4, lateral; 7, oral; both x 3. Jebel

| Thanais, basal 1 m of the Simsima Formation. 5, BMNH EE3572; apical detail, x 8. Jebel Buhays, section 1; loose in the scree derived
from the lowest 3 m of the Simsima Formatio

172

pentagonal caducous apical disc suggests that Plistophyma
belongs within the phymosomatoids.

Family STOMECHINIDAE Pomel, 1883
Genus CIRCOPELTIS Pomel, 1883

Pi isetiess—3 elle
fos 2hig1 37

Circopeltis? emiratus sp. nov.

Types. The holotype is BMNH EE3584 and paratypes are
BMNH EE3582, EE3583, EE3585 and EE3596.

OTHER MATERIAL. There is one other fragment tentatively
attributed to this species, BMNH EE3586.

OCCURRENCE. In the western Oman mountains this species

is found at the following levels:

Jebel Buhays section 1: loose in scree (derived from lower
beds of the Simsima Formation) (1).

Jebel Buhays section 3: immediately overlying prominent red
marly sand level (ca. 3 m above the base of the Simsima
Formation) (2).

Jebel Rawdah section 3: bed 6 (1). Section 3b basal 1 m (1).

Jebel Rawdah, section 4: bed 4 (1).

It is thus known from the Maastrichtian of the western Oman
Mountains.

DIAGNOSIS. Test low,. domal; apical disc hemicyclic with
plates firmly bound to the corona. Ambulacra straight,
plating compound in the phymosomatoid style with five or six
elements to a compound plate; pore-pairs arranged in arcs
from apex to peristome. There is a single large primary
tubercle to each plate with broad adradial and interradial
zones of secondary tuberculation.

DESCRIPTION. Tests range in diameter from 20-1 mm to 47-2
mm and in height from 11-8 to 23-5 mm (mean test height is
53% of test diameter). The test is circular in outline and
flattened ovate in profile, with the ambitus at approximately
40% of test height above the base.

The apical disc is hemicyclic, with the three anterior
oculars insert and the posterior two exsert (Fig. 37A). The
apical disc is 19-24% of the test diameter in diameter (mean
= 22%) and the periproct occupies approximately 55% of the
apical disc diameter. The periproct is D-shaped in outline,
with the slightly flattened edge abutting the madreporite.
Genital plate 2 is the largest and is tumid and covered in
dense madrepores. Other genital plates and all ocular plates
have a scattering of small miliary tubercles.

The ambulacra are 22-24% of the test diameter in width at
the ambitus. They are straight and taper gradually adapically.
All plates are compound in the phymosomatid style (Fig.
37B) and most are composed of five, or occasionally six,
elements. The three most adoral plates however are com-
posed of just four elements. Pore-pairs are arranged in arcs
and phyllodes are developed around the peristomial edge.
Each compound plate has a single large primary tubercle
(imperforate and non-crenulate) that overlaps all five (or six)
elements that make up the compound plate (Fig. 37C; Pl. 17,
fig. 1). There is a much smaller secondary tubercle lying
perradially, plus one or two scattered miliary granules. The
perradial zone is relatively broad. There are 14 compound
plates and 73 pore-pairs at 20 mm test diameter, rising to 18

A.B. SMITH

compound plates and 88 pore-pairs at 37 mm test diameter.
There are no sphaeridial pits.

Interambulacra are broad and carry two rows of primary
tubercles, set close to the adradial margin. There are 13
plates in a column at 20 mm test diameter, rising to 17 at 37
mm test diameter. Areoles are almost contiguous adorally
but are separated adapically. The mamelon is large and
imperforate and there is a narrow ledge that may retain faint
traces of crenulation in well-preserved individuals. Adradi-
ally there is a narrow band of small secondary tubercles and
miliary granules. Interradially the plates are slightly |
depressed and there are small and irregularly scattered sec- |
ondary tubercles throughout (PI. 17, fig. 2). Both columns
reach the peristomial margin.

The peristome is about 42% of the test diameter across. It
is slightly invaginated. Buccal notches are moderately well-
developed and have a thickened lip. Lantern and spines are
unknown.

REMARKS. This species has a phymosomatid style of ambu-
lacral compounding. However, the apical disc structure dif-
ferentiates it from the great majority of phymosomatids, since
these almost all have monocyclic apical discs that are typically
caducous. Only Glyptocidaris has a comparable apical disc.
Furthermore, this species has tuberculation that is virtually
non-crenulate, whereas other genera, including Glyptoci- |
daris, show stronger crenulation. The species is here tenta- |
tively assigned to the genus Circopeltis. Circopeltis has
polygeminate plate compounding and a hemicyclic apical
disc. Circopeltis also has non-crenulate tuberculation.. How-
ever, its ambulacral compounding style is unreported and it is
not yet known whether it is phymosomatoid.

It differs from Phymechinus? perplexus in its finer tubercu-
lation, with more extensive scattered secondary tubercula-

arcs throughout. In Phymechinus pore-pairs become irregu- |
larly biserial adapically. Furthermore, Circopeltis? has a |
larger peristome and much less well developed phyllodes than |
does Phymechinus? perplexus from the same levels. How-
ever, there is little doubt that the two forms are rather closely | |
related.

Genus PHYMECHINUS Desor, 1856

Phymechinus? perplexus sp. nov PI. 15, figs 4-10; Figs |
38, 308

Types. Holotype EE3579, paratypes BMNH EE3581, |

JB E25 oil, HSs5 98), BSI).

MATERIAL STUDIED. 14 specimens, of which biometric data | -
was derived from the following: BMNH EE3578-79, EE3581,
EE3989, EE3991-94, EE3619.

OCCURRENCE. All specimens come from Jebel Rawdah as
follows:

Section 1: bed 4 (2).

Section 2: bed 6 (2); bed 8 (4); bed 11 (1); loose in scree (4).
Section 4: bed 8/9 (1).

DIAGNOsIS. Apical disc small, caducous. Ambulacra
polygeminate with seven or eight elements to a compound
plate. Plate compounding phymosomatid-style. Pore-pairs
arcuate or irrgularly multiple above. One primary imperfo-
rate, crenulate tubercle on each ambulacral and interambu-

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 173

PLATE 15

figs 1-3 Circopeltis? emiratus sp. nov. BMNH EE3584, holotype; 1, apical; 2, oral; 3, lateral; all x 2. Jebel Buhays, section 3, ca. 3m

| above the base of the Simsima Formation.

jigs 4-10 Phymechinus? perplexus sp. nov. 4-6, BMNH EE3591, paratype; 4, oral; 5, apical; 6, lateral; all x 2. Jebel Rawdah, section 2,

| bed 8. 7, BMNH EE3579, holotype; lateral, x 1. Jebel Rawdah, section 2, bed 8. 8-10, BMNH EE3581, paratype; 8, lateral; 9, apical; 10,
| oral; all x 2. Jebel Rawdah, section 2, loose in scree derived from beds 3-10.

174

Ww
o

N
°o

Test height (mm)

fo)

Width at ambitus (mm)

20 30 40

Test diameter (mm)

=
[2)
fo)

Number of pore-pairs in column

20 30
Test diameter (mm)

Number of interamb. plates in column

30
Test diameter (mm)

Fig. 38 Biometric data for Phymechinus? perplexus sp. nov.

lacral plate. Mamelons very large and crenulation feebly
developed. Lower surface flat, peristome small with feeble
buccal notches. Phyllodes extremely well-developed.

DESCRIPTION. Tests are 11 to 60 mm in diameter and 5-8 to
31-2 mm in height (test height 50-60% of diameter; mean =
54%, SD = 3-2%, N = 9). They are circular in outline and
bun-shaped in profile, with a broad base and depressed
conical upper surface (Pl. 15, fig. 8). The ambitus lies at
approximately 30% of test height above the base.

The apical disc is small and circular, only 16-28% of the
test diameter across (PI. 15, fig. 9). It is proportionally
smaller in larger individuals (Fig. 38). No specimen retains
any apical disc plating, but to judge from the small size of the

Diameter (mm)

A.B. SMITH

8 Ambulacrum
@ Interambulacrum

Test diameter (mm)

®@ Peristome
® Apical disc

20 30 40 60

Test diameter (mm)

opening, plating is almost certain to have been dicyclic or
hemicyclic.

Ambulacra are only slightly narrower than interambulacra
at the ambitus and measure 23-28% of the test diameter in
width (mean = 25:5%, SD = 1:8%, N = 9). They are more
or less straight, tapering slightly adorally and more signifi-
cantly adapically. All plates are polygeminate with seven
(rarely six or eight) pore-pairs to an ambital plate. Plate
compounding is in the phymosomatid style (Figs 39A, B).
Pore-pairs are strongly arced at the ambitus, but tend to
become irregularly biserial or pleuriserial adapically and
adorally (Figs 38C, D). Close to the apex, pore-pairs once
again become uniserially arranged. Adapically, small second-
ary tubercles occur scattered within the pore zone. However,

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

there are a few specimens (e.g. EE3581) in which the
pore-pairs are in uniserial arcs right to the apex. Each plate
carries a single large primary tubercle that occupies most of
the plate (Pl. 15, fig. 7; Figs 38A, B). This has a relatively
well-developed imperforate mamelon and is distinctly crenu-
late, though the surrounding platform is not very broad.
Areoles are circular and separated by a single ring of small
secondary and miliary tubercles. There are 53 pore-pairs in a
column at 11 mm test diameter, rising to about 100 pore-pairs
at 34 mm test diameter (Fig. 38). Tubercles are largest at the
ambitus and decrease in size gradually both adapically and
adorally. Adorally there are very strong phyllodes composed
of circular pore-pairs with well-developed periporal muscle
attachment areas (PI. 15, fig. 10). These phyllodes are so
large as to make up more than half of the ambulacral width.

Interambulacra are relatively narrow, being only 30-37%

of the test diameter in width at the ambitus. There are 10
plates in a column at 11 mm test diameter, rising to 16 at 34
mm test diameter. Each plate carries a large primary tubercle
which is imperforate and weakly crenulate. In smaller speci-
mens (i.e. less than 35 mm) these are the only large tubercles
present, although a distinct small secondary tubercle is
present both adradially and interradially. In specimens
around 58 mm test diameter the adradial secondary tubercle
reaches approximately half the diameter of the primary
tubercle at the ambitus and adorally, and there are additional
smaller secondary tubercles close to the adradial suture.
Primary and secondary tubercles extend to the apex and

_peristome. The primary tubercles are non-confluent, being
separated by a single row of miliaries.

The peristome is 30-55% of the test diameter in width
(proportionally smaller in larger individuals) and is hardly
invaginated, the entire lower surface being flat. Buccal
' notches are small and poorly developed. Spines, lantern and
_ pedicellariae are all unknown.

| REMARKS. It is with some slight hesitation that I refer the
_ new species P? perplexus to this genus. This species has stout
'tubercles with large mamelons and virtually no platform.
‘Nevertheless fine crenulation is developed around the
mamelon, though it is often difficult to see unless preserva-
tion is near perfect. The ambulacral plate compounding is

‘Fig. 39 Camera lucida drawings of plating in Phymechinus?
perplexus sp. nov., BMNH EE3581. A, ambital ambulacral plate;
B, adapical ambulacral plate; C, adapical pore-pair arrangement;
D, adoral pore-pair arrangement. Scale bar = 1 mm.

phymosomatid in style, with adapical pore-pairs either
strongly arcuate or, more often, actually multiple in an
irregular way. The apical disc is small and unlikely to be
monocyclic, but plating is not preserved. The type species of
Phymechinus, P. mirabilis (Agassiz) comes from the Middle
Oxfordian and has a similar overall shape, small apical disc
and stout tuberculation. Well-preserved specimens (e.g. Hess
1975, pl. 37, fig. 5) apparently show feeble crenulation.
Unfortunately, none of the specimens I have examined show
the style of ambulacral compounding. P? perplexus differs
from the type species in having a proportionally smaller
peristome and better developed phyllodes. It also has much
less well-developed buccal notches. In tuberculation style it is
very similar to Schluter’s species Phymechinus cretaceus from
the B. mucronata Zone, Upper Campanian of Ciply, Bel-
gium, but this species has a much larger peristome and much
less well-developed phyllodes. This species is easily distin-
guished from Phymosoma cf. paroni Checchia-Rispoli by its
very much smaller apical disc, more subconical profile and
compound amulacral plates that incorporate more than five
elements. It is also easily distinguished from Actinophyma.
Although Actinophyma has a similar arrangement of pore-
pairs adapically, forming rather irregular multiple columns
with intersperced tubercles, it is very different adorally. The
peristome in Actinophyma is invaginated and the pore-pairs
remain uniserial and rather widely spaced across the entire
oral surface. In P? perplexus the pore-pairs form a very
strong phyllode and the oral area is broad and flat.

It is distinguished from Circopeltis emiratus by its very
much coarser tuberculation, smaller peristome, more
polygeminate ambulacral compounding and cadoucous apical
disc. Furthermore, C. emiratus never developed biserial
pore-pairs adapically.

Genus ECHINOTIARA Pomel, 1883

Echinotiara perebaskinei Lambert, 1930 Pl. 16, figs
1-6; Pl. 17, figs 3, 6, 7; Figs 40, 41

1930 Echinotiara perebaskinei Lambert, in Lambert & Pere-
baskine: 472, pl. 38, figs 1-5.

Types. The two specimens figured and described by Lambert
and presumably in the Lambert Collection, Université de
Paris VI, France.

MATERIAL STUDIED. 82 specimens of which only the follow-
ing were used in the biometric analysis: BMNH EE3756-57,
EE3761-63, EE3767-69, EE3772, EE3774, EE3782,
EE3785-88.

OCCURRENCE. Along the western margins of the Oman

mountains this species is confined to the lowest arenaceous

levels of the Simsima Formation at Jebel Rawdah. It occurs

as follows:

Jebel Rawdah, section 2: bed 4 (6); bed 6 (9); bed 8 (49); bed
11 (3); loose in lower scree (17).

Jebel Rawdah, section 3b: bed 3 (1).

Jebel Rawdah, section 4: bed 2 (1); bed 8/9 (1).

The species was originally described from the ‘Calcaires
inferieur a Libycoceras, Maastrichtien’ at Oued Tarinkat,
Tchi-Dermine and Oued Tinamassine in tke district of Gao,

176

Test height (mm)

Test diameter (mm)

& Amulacral pore-pairs
@ Interambulacral plates

Number in column

N
o

20 30
Test diameter (mm)

Fig. 40 Biometric data for Echinotiara perebaskinei Lambert.

Niger (Lambert & Perebaskine 1930). Amard ef al. (1981:
124) reported an Echinotiara cf. perebaskinei Lambert from
the Upper Maastrichtian of Tademait, Algeria, but as no
figures or description are given it is not possible to confirm
this record.

DiAGnosis. An Echinotiara with a relatively small caducous
apical disc, ambulacral pore-pairs in arcs of three, with very
strong phyllodes developed adorally. Interambulacral plates
with a single large primary tubercle and well-developed
adradial and interradial secondary tubercles that diminish in
size adapically.

DESCRIPTION. Tests range from 9 to 32 mm in diameter and
are more or less circular in outline. Test height is 40-52% of
test diameter (mean = 46%, SD = 3:7%, N = 14). In profile
the test is low conical (Pl. 16, fig. 3), with the ambitus
approximately 40% above the base.

The apical disc is always missing and plating can only have
been loosely fixed to the corona. The apical disc outline is
irregularly circular (Pl. 16, fig. 1) and is 21-29% of the test
diameter in length (mean = 24%, SD = 2:1%, N = 13). To
judge from the size, it must have been dicyclic or hemicyclic.

Ambulacra are trigeminate throughout with pore-pairs
arranged in distinct arcs of three (Fig. 41A). Ambulacra are
relatively broad, being approximatley 22% of the test diam-
eter in width at the ambitus. They taper gradually adapically,
but primary tubercles continue to the apex (Pl. 17, fig. 3).
Adorally the ambulacra remain broad and there are large,
well-developed phyllodes, which in specimens 25 mm in
diameter include at least 22 pore-pairs in each column (PI. 16,

Diameter (mm)

Number in column

® Peristome
@ Apical disc

Test diameter (mm)

® Amulacral pore-pairs
¢@ |Interambulacral plates

BS
lo)

20
Test diameter (mm)

30 40

fig. 5). Each compound plate carries a single large imperfo-

rate and non-crenulate tubercle that overlaps all three ele- |
ments (Fig. 41). Plate compounding is diadematoid in style,
with all three elements reaching the perradius and the middle |

plate pinched towards the centre of the plate (Fig. 41C). The
lowest element in each compound plate carries a small |
secondary tubercle adradially and perradially. There are 32 |

pore-pairs in a column at 9 mm test diameter, rising to 71 at |
32 mm test diameter (Fig. 40).

At the ambitus the interambulacra are 35-38% of the test |

diameter in width. Both columns reach the peristomial bor- |

der, although the interambulacra taper considerably towards |

the peristome. Plates at the ambitus are longer than wide and
are slightly curved. Each plate carries a single large imperfo- |
rate and apparently non-crenulate tubercle at its centre (PI.
17, fig. 7). The mamelon is slightly undercut but there is little |
surrounding platform developed. On the adradial side there
are one or two smaller secondary tubercles, while on the

interradial edge there is a single secondary tubercle (Fig. |
41C). These primary and secondary tubercles are relatively |

coarse and occupy most of the available space. There are, |
however, miliary granules and small tertiary tubercles along |
the adapical margin and in spaces adjacent to the primary
tubercle. There are 10 plates in a column at 9 mm test
diameter, rising to 16 at 32 mm test diameter. Primary
tubercles continue to the apex and there is no median naked
zone.

The peristome is circular and measures 35-50% of the test
diameter in diameter (it is proportionally larger in smaller
individuals (Pl. 16, figs 4, 5). It is only slightly sunken and

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 177

|PLATE 16

(Figs 1-6 = Echinotiara perebaskinei Lambert. Jebel Rawdah, section 2, beds 6-8. 1, 2, BMNH EE3756; 1, apical, 2, oral; both x 3. 3, 4,
BMNH EE3768; 3, lateral; 4, oral; both x 3. 5, 6, BMNH EE3788; 5, oral; 6, lateral; both x 3.

178

Fig. 41 Camera lucida drawings of plating in Echinotiara
perebaskinei Lambert. A, BMNH EE3769, ambulacrum from
apex (top) to peristome margin (bottom); B, BMNH EE3763,
ambulacrum from apex (top) to peristome margin (bottom); C,
ambital ambulacral and interambulacral plate, BMNH EE3763.
Scale bar = 1 mm.

buccal notches are small but distinct.

The perignathic girdle structure is seen in BMNH EE3783
and EE3788. It consists of two long peg-like auricles that do
not meet above the perradius. Spines and lantern are
unknown.

REMARKS. This. species lacks crenulate tuberculation,
although preservation is usually inadequate to be certain for
most specimens. It could easily be mistaken for Orthopsis
miliaris on account of its very similar tuberculation and test
shape. However, Orthopsis has perforate tuberculation and
its ambulacral pore-pairs are strictly uniserial, not arranged in
arcs of three as in Echinotiara. Furthermore, Echinotiara has
well-developed phyllodes that are never seen in Orthopsis.

The type material described and illustrated by Lambert
differs in apparently having slightly less well developed
phyllodes adorally at comparable sizes, but for the present
the two populations are treated as conspecific.

A.B. SMITH

Cohort IRREGULARIA Latreille, 1825

Order HOLECTYPOIDA Duncan, 1889
Family HOLECTYPIDAE Lambert, 1899
Genus COENHOLECTYPUS Pomel, 1883

Coenholectypus inflatus (Cotteau & Gauthier, 1895)
Pl. 18, figs 7-11; Figs 42, 43A, C

Holectypus inflatus Cotteau & Gauthier: 73, pl. 12,
figs 14.

21989 Holectypus (Caenholectypus) inflatus Cotteau &
Gauthier; Ali: 401, fig. 4 (1).

Types. The specimen described and illustrated by Cotteau &
Gauthier, from the late Cretaceous of Aftab, southern Iran.

1895

MATERIAL STUDIED. Thirty specimens, of which 16 were |

used in the biometric analysis (BMNH E82644, EE3399,
EE3401-04, EE3406-07, EE3409, EE3411-15, EE3417,
EE3429).

OCCURRENCE. There are two morphologies found along the
western foothills of the Oman Mountains:

Depressed variety: this occurs at Jebel Huwayyah, from an
uncertain horizon. It is also found at Jebel Buhays (sections 1
and 2) in the lowest limestones. It is more common at Jebel
Rawdah: in section 3 it occurs reasonably abundantly in bed 6
(6) and is found at a comparable level in section 4 (beds 19
and 21—22 and at top of measured section) (10). It also occurs
frequently at and immediately above the level of the first
major red parting in section 3 (bed 5).

More inflated, rounded forms are recorded from Jebel
Huwayyah (unknown horizon); and from Jebel Rawdah,
Section 2, bed 6, section 3, bed 10, section 4, bed 13.

DESCRIPTION. Tests are circular in outline and range from
around 13 mm up to 56 mm in diameter. In profile their
height varies from low conical to almost subglobular. Test
height is 47-77% of test length (Fig. 42). The ambitus is well
rounded and lies a little below midheight (Pl. 18, fig. 11).

The apical disc is small and compact with five gonopores.
Genital plates are small and pentagonal and separated from
one another by ocular plates that are almost as large (Fig.
43C). Madrepores occupy the entire central region.

Ambulacra are simple and uniserial, except adapically in
larger specimens (40 mm diameter plus), where pore-pairs
become slightly offset creating an incipient biserial arrange-
ment. There is no pore crowding towards the peristome (Fig.
43A) and all plating is simple. Only close to the peristome
does ambulacral plating become differentiated into triads
with every third plate becoming enlarged. All pore-pairs are
nonconjugate. There are approximately 100 pore-pairs in a
column at 18 mm test length, rising to 185 at 56 mm test
length (Fig. 42).

Interambulacra are standard in their structure. The
periproct is oval in outline and lies close to the peristome and
well separated from the ambitus (PI. 18, fig. 7; Fig. 43A). It
opens between interambulacral plates 2a,b and 6b,7a or 7a,b.
Its width is 55-81% of its length (mean = 65%, SD = 7%, N
= 17). The distance separating the periproct and peristome is
small, only 20-50% of the periproct length; whereas the
distance separating the periproct from the ambitus is much
greater, being 110-260% of the periproct length (greatest in
largest individuals).

The peristome is circular with feeble buccal notches. It is
15-35% of test length in diameter (mean = 23%, SD =

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 179

‘PLATE 17
Figs 1,2 Circopeltis? emiratus sp. nov. BMNH EE3584, holotype, details of ambital tuberculation; 1, ambulacrum, x 1325

| interambulacrum, x 5.

Figs 3, 6,7 Echinotiara perebaskinei Lambert. 3, 6, BMNH EE3774; 3, apical, x 6; 6, ambital detail, x 5. Jebel Rawdah, section 2, bed 4.
| 7, BMNH EE3785, lateral detail, x 6. Jebel Rawdah, section 2, beds 6-8.
Figs 4,5 Salenia nutrix Peron & Gauthier. 4, BMNH EE3627, apical, x 5. Jebel Rawdah, section 1, bed 3. 5, BMNH EE3634, lateral, x 5.
Jebel Rawdah, section 2, loose in scree at level of bed 11.

180

Fig. 42

4-6%,

40 é

@ C. cf baluchistanensis
@ C. inflatus e

ao)

220

F 10

20 30 40 50 60

Test diameter (mm)

C. cf. baluchistanensis
C. inflatus

w
fo}

N
(o}

Number of interamb. plates in column

rS)

20 30 40

Test diameter (mm)

=
lo}
a

C. cf baluchistanensis
C. inflatus

fee)

Periproct length (mm)
o

2
10 20 30 40 50 60
Test diameter (mm)
10r w C cf. baluchistanensis

@ C. inflatus

Periproct width (mm)

Periproct length (mm)

Biometric data for Coenholectypus inflatus (Cotteau & Gauthier) and C. cf. baluchistanensis (Noetling).

N = 16), being relatively larger in small individuals.

REMARKS. This species is easily recognised by its rounded
profile, and its small periproct that lies close to the peristome
and well separated from the ambitus. Although Cotteau &
Gauthier’s (1895) description is sketchy, it is readily recognis-

Distance between peristome

and periproct (mm)

Distance from periproct

to ambitus (mm)

able from the illustrations given. Ali (1989) described two)
species of Coenholectypus from the western Oman mountains|

¢
a

Number of pore-pairs in column

A.B. SMITH

200

BS C. cf baluchistanensis
@ C. inflatus ey

180

160

140

120

100

30 40 60

Test diameter (mm)

10)

2 4 6 8
Test length (mm)

Peristome length (mm)

20 30 40

Test diameter (mm)

SO 60

Test length (mm)

. inflatus and C. larteti (Cotteau). Unfortunately the latter is

Cenomanian species (Smith et al. 1990). Because Ali gave! —
no description and only illustrated the aboral surfaces of his

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 181

Pie yee
og
SG

| PLATE 18

)Figs 1-6 Coenholectypus cf. baluchistanensis (Noetling). 1-3, BMNH EE3389; 1, apical; 2, oral; 3, lateral; all x 2. Jebel Buhays, section 1;
loose in the scree derived from the lowest 3 m of the Simsima Formation. 4-6, BMNH EE 3395; 4, apical; 5, oral; 6, lateral; all « 2. Jebel
Rawdah, section 2, bed 11.

Figs 7-11 Coenholectypus inflatus Cotteau & Gauthier. 7, BMNH EE4330; oral, x 1. Jebel Buhays, section 1; loose in the scree derived

| from the lowest 3 m of the Simsima Formation. 8, 9, BMNH EE3412; 8, lateral; 9, oral; both x 2. Jebel Buhays, section 2, lowest part of

| the Simsima Formation. 10, 11, BMNH EE3424; 10, apical; 11, lateral; both x 1-5. Jebel Rawdah, section 3b, loose, from higher part of

the section.

182

two forms, it is impossible to tell to which species either
belongs. His assignment of one to C. inflatus is therefore
accepted tentatively.

C. inflatus differs from C. cf. baluchistanensis (Noetling),
the other species described here, on several counts. Firstly, it
has finer tuberculation on its oral surface at comparable sizes.
Secondly, its peristome is proportionally smaller at compa-
rable sizes, and thirdly, its periproct is smaller and more
removed from the ambitus at all sizes (Fig. 42).

There is a stratigraphical variation in profile. The taller
forms, exactly comparable in form with the Iranian type, are
found lower down in the sections in the coarse calcarenites: in
bed 2 at Jebel Rawdah section 4, In bed 6 at section 2, and in
bed 13 at section 4. The more depressed forms are found
higher up in the more orbitoline-rich limestones and may
represent deeper water morphological varieties.

Coenholectypus cf. baluchistanensis (Noetling, 1897)
Pl. 18, figs 1-6; Figs 42, 43B, D

cf. 1897 Holectypus baluchistanensis Noetling: 18, pl. 3, fig.
3.

MATERIAL. Thirteen specimens (BMNH EE3386-98).

OCCURRENCE. In the western Oman mountains this species
was found at the following localities and horizons:

Jebel Buhays, section 1: loose in scree (3) and bed 8 (1).
Jebel Thanais, lowest 2 m of limestone section (1).

Jebel Rawdah, section 1: 20 cm below the top of bed 5 (1).
Jebel Rawdah, section 2: bed 11 (9).

Fig. 43

Camera lucida drawings of plating in Coenholectypus. A, C, C. inflatus (Cotteau & Gauthier), BMNH EE3412: A, oral plating; C,

A.B. SMITH

It was originally described from the Maastrichtian of Bal-
uchistan.

DESCRIPTION. Tests are circular in outline and range from
18-37 mm in diameter. They are moderately inflated in
profile (Pl. 18, fig. 3), with a test height 55-62% of test
length. The ambitus is rounded and lies a little below
mid-height.

The apical disc has five gonopores. The four genital plates
are small and pentagonal and separated from each other by
ocular plates that are similar in size (Fig. 43D). The
madreporite plate is large and often tumid. Madrepores are
well developed over the central area.

Ambulacra are uniserial throughout, with simple noncon-
jugate pores. There is no hint of incipient biseriality adapi-

cally, nor any pore crowding adorally. All plates are simple, |

with triad development only appearing towards the peris-
tome.

The interambulacra are standard in structure. The
periproct is relatively large and opens between interambulac-
ral plates 2a,b and 7a,b. It is oval in outline, pointed at both
ends. Its width is 60-76% of its length (mean = 70%, SD =
5:7%, N = 7). It opens close to the peristome (PI. 18, fig. 2;
Fig. 43B), separated by 20-45% of the periproct length from
the peristome (mean = 32%, SD = 8-4%, N = 7). It is
separated from the ambitus by only a small distance in
smaller individuals, some 28% of periproct length at 17 mm
test length, but this increases in larger individuals to reach
65% of periproct length at a test length of 35-37 mm.

The peristome is circular in outline with deep and well
marked buccal notches. It is 10-22-5% of test length in

apical disc. B, D, C. cf. baluchistanensis (Noetling), BMNH EE3389: B, oral plating; D, apical disc. Scale bars: A, B = 5 mm; C, D = 1

mm.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

diameter (being relatively larger in small individuals).

REMARKS. This species is easily distinguished from the other
Coenholectypus that occurs here, C. inflatus, by its larger
peristome, larger periproct that extends much closer to the
ambitus, and by its coarser adoral tuberculation. It also has
deeper buccal notches.

C. baluchistanensis Noetling, from the Maastrichtian of
Baluchistan, was established on the basis of two specimens 31
and 38 mm in diameter. Like the species described here, this
has a relatively large peristome occupying much of the lower
surface. Unlike the arabian specimens, the periproct of the
Baluchistan species extends closer to the ambitus in the
illustrated specimen. However, Noetling states that all speci-
mens are crushed to some degree, and the illustrated speci-
men is damaged posteriorly. Therefore I suspect that the
periproct is in reality slightly more distant from the ambitus
than is actually shown. Until new material of C. baluchistan-
ensis is available, the identification of Omani material as this
species must remain tentative.

C. subcrassus Peron & Gauthier, from the early Maastrich-
tian of Algeria and Tunisia, differs from C. baluchistanensis

_ in having a much larger periproct that occupies virtually the
» entire oral surface of the posterior interambulacrum. It is
illustrated as having a broad, flat, adambital margin, unlike
{ the arabian species. However, C. subcrassus and C. baluchis-

' tanensis are sister taxa.

Genus COPTODISCUS Cotteau & Gauthier, 1895

| Coptodiscus magniproctus sp. nov. PI. 19, figs 4-7; Figs
44, 45

BMNH

| Types. Holotype, BMNH_ EE3716,
_ EE3715. There are no other specimens.

paratype,

OCCURRENCE. Both specimens come from the base of the
“silty Loftusia beds (bed 1), at Jebel Huwayyah, section 2.

| DIAGNOsIs. A Coptodiscus with a relatively large periproct
occupying most of the oral surface of the posterior interam-
bulacrum and opening between interambulacral plates 2 and
7. Aboral interambulacral ornament comprising a series of
fine sutural pits and a set of pits along the midline of the each
| plate.

DESCRIPTION. Both specimens are small, but have open
‘gonopores and are thus mature individuals. The smaller
‘specimen is 10 mm in diameter, the larger 16 mm. Both are
‘circular and low conical in profile, with the ambitus posi-
\tioned relatively low down. Test height is about 40% of test
‘diameter.
The apical disc is small and compact (Fig. 45A). There are
five genital plates, each perforated by a gonopore. The ocular
plates are almost as large as the genital plates, but are exsert
and only oculars III and II abut the madreporite plate. The
Zenital plates are contiguous around the posterior and lateral
\margins of the madreporite.
| Ambulacra are uniserial and simple throughout, pores
secoming slightly more widely spaced towards the peristome.
There are three and a half ambulacral plates to an ambital
mterambulacral plate.

The peristome is relatively large, 25% of test diameter in
liameter. It is not much invaginated. The periproct is rela-
‘ively large, being 3-6 mm in length (22% of test diameter) by

@C. magniproctus

Periproct width (mm)

“10 20 30
Test diameter (mm)

2 @C. magniproctus

Le,)

° 3

a
2

10 20 30
Test diameter (mm)

Fig. 44 Biometric data for species of Coptodiscus. Data from the
type series of Coptodiscus noemiae Cotteau & Gauthier. The
holotype of C. magniproctus sp. nov. is also plotted.

A B
O
fe
a8 COS

Fig. 45 Camera lucida drawings of Coptodiscus magniproctus sp.
nov., BMNH EE3716. A, Apical disc; B, ambital interambulacral
plate, adradial margin to right. Scale bar = 1 mm.

2:3 mm. It is pointed at both ends and lies close to both the
peristome and the ambitus. Only 1-3 mm separates the
periproct from the peristome and 1-2 mm separates the
periproct from the ambitus. The periproct opens at interam-
bulacral plate 2 and extends to interambulacral plate 7 (Fig.
44).

Tuberculation is standard with three or four primary
tubercles on an ambital interambulacral plate. The surface of
the test is ornamented by rows of fine pits (PI. 19, fig. 4; Fig.
45B). These are arranged along the horizontal sutures, but
are also developed along the midline of the plate in between
the primary tubercles.

REMARKS. Amongst holectypoids, ornamentation of the
test, as seen in this species, is found only in the genus
Coptodiscus. Only one late Cretaceous species of Coptodis-
cus has ever been described, C. nomiae Cotteau & Gauthier
(1895) (Pl. 19, figs 1-3; see Kier 1972 for a detailed descrip-
tion). C. nomiae, which comes from the ‘Senonian’ of south-

184 A.B. SMITH |

PLATE 19

Figs 1-3 Coptodiscus noemiae Cotteau & Gauthier. Syntype, from the Morgan Collection, Museum d’Histoire Naturelle, Paris; 1, apical, x
5; 2, apical, x 2; 3, oral, x 2; Senonian, Khianan, Iran. }

Figs 4-7 Coptodiscus magniproctus sp. nov. Jebel Huwayyah, section 2, bed 1. 4, 6, 7, BMNH EE3716, holotype; 4, detail of adapical
ornamentation, x 10; 6, oral, X 3; 7, apical, x 3. 5, BMNH EE3715, paratype, apical surface, x 5.

Figs 8-11 Conulus douvillei Cotteau & Gauthier. BMNH EE4306; 8, apical; 9, oral; 10, lateral; 11, posterior; all x 2. Jebel Thanais, lowest
2 m of the Simsima Formation. }

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 185

40
8 G. bleicheri
@ C. douvillei =
x E
E 30 E
E *
y a
[>] =
3 20 2
5 E
” ©
is E
10 2
LJ
i=
©
a
(0)
(0) 10 20 30. 40 50 2 3 4 5 6 U/ 8
AAG Test diameter (mm) 5a Peristome major axis (mm)
8 G. Dleicheri 4 S G. bleicheri
© C. dowvillei on 50 © C. dowillei
40
5 -_-_
5 E
° ~ 30
: g
a AS)
= 2 20
3 3
NM =
é 10

10) 10 20 30 40 50
Test diameter (mm) Test length (mm)

14
8 G. bleicheri

12 @ C. douvillei a

Height to base of periproct (mm)

Test diameter (mm)

Fig. 46 Biometric data for ‘Globator’ bleicheri (Thomas & Gauthier) and Conulus douvillei (Cotteau & Gauthier).

ern Iran and the late Campanian of Saudi Arabia and Oman, Family CONULIDAE Lambert, 1911
differs from our species in several important details. Firstly, Genus ‘GLOBATOR’ Agassiz, 1840
its periproct is very much smaller at comparable sizes (Fig.
|44) and it lies well separated from the peristome, with at least
three and usually four interambulacral plates in each column
Separating the two openings, as opposed to the two in C.
magniproctus. The periproct is also considerably more
rounded in C. nomiae. A second immediately apparent
difference is in the style of ornamentation developed abo-

REMARKS. The genus Globator was erected by Agassiz
(1840) for small, ovoid conulids with large periprocts opening
above the ambitus. The type species is Globator nucleus
Agassiz, but this is based on a juvenile Conulus rotundus
Goldfuss and thus Globator falls into synonymy with Conu-
lus. However, there is a distinct clade of ovoid conulids with
2 . 3 ; large, supra-ambital periprocts that can be recognized. This
rally. In C. nomiae there is a single laterally extensive sutural one is ie a. Pare! (Smith & Wright, in prep.)

pit on either side of the primary tubercle, also, intraplate pits ee
| : : 5 and for the moment we retain its members under the name
are not as well developed, whereas in C. magniprocta there is ‘Globator’

4 well-developed row of sutural pits (compare PI. 19, figs 1,

4),

}

186

‘Globator’ bleicheri (Gauthier, 1889) PI. 20, figs 1-10;
Figs 46, 47A, B, F-I, 48A

1889 Pyrina bleicheri Gauthier: 51, pl. 3, figs 15-18.

1895 Pyrina orientalis Cotteau & Gauthier: 68, pl. 11, figs
1-8.

1897 Pyrina zumoffeni de Loriol: 158, pl. 7, fig. 1.

1967 Pyrina ovulum Agassiz; Devries: 177, pl. 5, figs 19-21.

1987 Pseudopyrina bleicheri (Thomas & Gauthier);
Zhagbib-Turki: 167.

1989 Globator orientalis (Cotteau & Gauthier); Ali: 403,
fig. 5 (4-5).

MATERIAL STUDIED. This is a common species in the lower
beds of the Simsima Formation and a large number of
specimens were available for study. There are 208 specimens
in the collections made. Of these the following were mea-
sured: BMNH EE4107, EE4116-19, EE4121, EE4129,
EE4130, EE4132-34, EE4139-41, EE4144-45, EE4154-57,
EE4163, EE4172, EE4174, EE4186, EE4208, EE4217,
EE4232, EE4249, EE4251, EE4253-54, EE4257.

OCCURRENCE. The species was first described from the late

Cretaceous (late Campanian) of Jebel Atra, Tunisia. It has

also been described from the Upper Senonian of Derre-i-

Chahr and Endjir-kouh, southern Iran (Cotteau & Gauthier

1895) and the late Cretaceous of Palestine (de Loriol 1897)

and Turkey (Devries 1967). In the western margins of the

central Oman Mountains this species is found at the following

localities and levels:

Jebel Buhays, section 1: loose in scree (96).

Jebel Buhays, section 2: loose in scree (3).

Jebel Buhays, section 3: lowest 2 metres (1).

Jebel Thanais: lowest couple of metres of section (5).

Jebel Agabah: basal shell bed (2).

Jebel Faiyah, section 1: arbaciid level ca. 4 m above base
(12).

Jebel Huwayyah, section 1: beds 14 and 15 (54).

Fig. 47 Camera lucida drawings of plating in ‘Globator’ and Conulus. A, B, F-I, ‘G.’ bleicheri (Thomas & Gauthier). A, B, BMNH EE4187;
A, adapical ambulacral plating; B, adoral ambulacral plating, peristomial margin at base; F, BMNH EE4186, apical disc; G, BMNH
EE4148, apical disc; H, BMNH EE4154, apical disc; I BMNH EE4151, peristomial plating. C-E, Conulus douvillei (Cotteau & Gauthier),
apical disc plating: C, BMNH EE4204; D, BMNH EE4277; E, BMNH EE4211. Scale bars = 1 mm.

A.B. SMITH

Jebel Rawdah, section 1: bed 6 (1); top of bed 4 (15); bed 3
(49); loose (2).

Jebel Rawdah, section 2: bed 6 (1); bed 11 (10); bed 13 (1);
bed 14 (7); bed 16 (3); bed 19 (8); bed 20 (1); bed 21 (1) |
loose in scree just above bed 12 (2); loose in scree, mostly |
near base (14).

Jebel Rawdah, section 3: bed 1 (11); bed 5 (9); bed 9 (1); bed
11 (1); loose in scree (3).

Jebel Rawdah, section 4: bed 2 (9); bed 4 (4); bed 5 (1); bed 8
(3); bed 10 (10); bed 14 (3); bed 20 (1).

DIAGNOsIS. An oval, rather depressed ‘Globator’ with a |
large, strongly ellipsoidal peristome. The periproct lies high |
on the posterior surface and is visible from above but not | |
from below. Pore-pairs in weak arcs only towards the peris- |
tome, not forming multiple rows. Genital plates 3 and 4]
separated by ocular plate IV in adults.

DESCRIPTION. Tests range from 9 to 35 mm in length and are |
oval in outline and profile. Test width is 82-91% of test
length (mean = 86%, SD = 2:6%, N = 32; Fig. 46) with the |
widest point on the test coincidental with the posterior
portion of the anterior ambulacra. Test height is 61-80% of |
test length (mean = 70%, SD = 5:3%, N = 32) and the tallest |
point on the test is subcentral. Tests in profile have a |}
relatively broad, flat apex and base and a rounded ambitus | —
(Pl. 20, figs 3, 8).

The apical disc is more or less central and is tetrabasal (Fig. |
47F-H). Genital plate 2 is considerably larger than the other | _
four genital plates and is covered in madrepores. Genital Fi
plate 3 is the smallest and in the great majority of specimens .
is separated from genital plate 4 by ocular plate 4, which | —
abuts genital plate 2. Genital plates 3 and 4 are found in tl
contact only in small individuals. The posterior pair of genital | )
plates are in contact posterior to genital plate 2. Ocular plates | |
are pentagonal in outline and project. ti

Ambulacra are uniserial and pore-pairs are undifferenti- | |
ated. Above the ambitus they are very strictly uniserial (Fig. |

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 187

sei)
OY

3 ae
S209
SSo 0

£95
ot

LATE 20

igs 1-10 ‘Globator’ bleicheri (Peron & Gauthier). 1-4, Topotype specimen of Pyrina orientalis Cotteau & Gauthier, from the Morgan
Collection, Museum d’Histoire Naturelle, Paris; 1, oral; 2, apical; 3, lateral; 4, posterior; all x 2. Senonian, Poucht-e-Kouk, Iran. 5, 6, 10,
BMNH EE4251; 5, apical; 6, oral; 10, posterior; all x 2. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the

| Simsima Formation. 7-9, BMNH EE4208; 7, posterior; 8, lateral; 9, apical; all x 2. Jebel Buhays, section 1; loose in the scree derived from

' the lowest 3 m of the Simsima Formation.

igs 11-16 Conulus douvillei (Cotteau & Gauthier). 11, 12, topotype specimen from the Morgan collection, Museum d’Histoire Naturelle,

| Paris; 11, oral; 12, lateral; both x 2. Senonian, Khianan, Iran. 13-16, BMNH EE4308; 13, oral; 14, apical; 15, posterior; 16, lateral; all
| 2. Jebel Thanais, lowest 2 m of the Simsima Formation.

188

Fig. 48 Camera lucida drawings of adoral pore arrangement. A,
‘Globator’ bleicheri (Thomas & Gauthier), BMNH EE4187; B,
Conulus douvillei (Cotteau & Gauthier), BMNH EE4309. Scale
bar = 1 mm.

47A), but towards the peristome they become weakly arcuate
and reduce in pore-diameter size (Figs 47B, 48A). There are
about 88 pore-pairs in a column at 18 mm test length, rising to
127 at 32 mm test length (Fig. 46). Plates are compound in the
pyrinoid style, with a single small demiplate in each triad
(Figs 47A, B). All plate sutures are denticulate.

The periproct is large and tear-drop shaped, pointed adapi-
cally (Pl. 20, figs 7, 10). It lies on the posterior surface
relatively high on the test, so that it is visible when viewed
from above, but not from beiow. Periproct height is 28-46%
of the test height (mean = 36%, SD = 4:6%, N = 29) and
periproct width is 56-80% of its height (mean = 65%, SD =
5:8% , N = 27). The distance from the base of the periproct to
the base of the test is 38-62% of the test height (mean =
49%, SD = 5-6%, N = 31).

The peristome is oblique and broadly fusiform in outline,
with the long axis running from interambulacrum 3 to ambu-
lacrum I (PI. 20, fig. 6). There is hardly any invaginated lip

A.B. SMITH
|

developed around the peristome, although the oral surface !
does curve inwards towards the periproct slightly. |

Tuberculation is uniform throughout, with semi-regular
and slightly sunken primary tubercles scattered over the)
surface, surrounded by a very dense miliary granulation. |
There is no internal butressing.

REMARKS. This species is easily distinguished from the other —
species of Conulidae, Conulus douvillei, that occurs here. C. |
douvillei has a periproctal opening that lies close to the base},
of the test, whereas the periproct in ‘G.’ bleicheri lies high on!
the posterior and is separated by a considerable distance from
the base. This is not a size-related character since there is
clear separation of the two species at all sizes (Fig. 46). A’
second difference concerns the development of phyllodes
adorally. In ‘G.’ bleicheri the pore-pairs become slightly)
arcuate adorally (Fig. 48A), but even in the largest specimens,
they never become triserially arranged. Adoral pore-pairs in’
C. douvillei, by contrast, are arranged triserially across much
of the oral surface (Fig. 48B). Finally, in the apical disc’
plating of ‘G.’ bleicheri genital plate 2 almost always reaches
to ocular IV separating genital plates 3 and 4. In C. douvillei
genital plate 2 does not reach ocular plate IV and genital
plates 3 and 4 maintain firm contact. :

This species has previously been recorded from the Oman)
mountain region by Ali (1989) and Smith (in Skelton et al.
1990) under the name Globator orientalis (Cotteau &
Gauthier). ‘G.’ orientalis (P1. 20, figs 1-4) was described from)
the late Cretaceous of southern Iran by Cotteau & Gauthier
(1895). However, it appears virtually indistinguishable in
form to ‘Globator’ bleicheri Thomas & Gauthier, from the)
late Campanian of Tunisia (Zhagbib-Turki 1987). The only
slight difference between these two forms is that “G.’ orienta-
lis may have a slightly smaller peristome. For the present,|
however, the two species are synonymized.

Genus CONULUS Leske, 1778 |

Conulus douvillei (Cotteau & Gauthier, 1895) PI. 19,
figs 8-11; Pl. 20, figs 11-16; Figs 46, 47C-E, 48B

1895 Echinoconus douvillei Cotteau & Gauthier: 70, pl. 11,

figs 9-13.

1932 Pyrina mortenseni Checchia-Rispoli: 21, pl. 2, figs 1-3,
le 3h, whys I 2.

21967 Conulus douvillei (Cotteau & Gauthier); Devries:

184, pl. 5, figs 22-25.
1972 Globator mortenseni (Checchia-Rispoli); Kier: 70, figs
35, 36, pl. 44, figs 1-7.
1989 Globator mortenseni (Checchia-Rispoli); Ali: 403, Fig.
4 (6-7).

TypPES. The syntypes are the five specimens whose dimen-
sions are cited by Cotteau & Gauthier (1895: 70). They may
be represented amongst material in either the Cotteau Col-|
lection (Lyon) or the Morgan Collection (Museum d’Histoire
Naturelle, Paris), but none have been definitely identified.

MATERIAL STUDIED. This species is less common than “Glo-
bator’ bleicheri, but is never the less well represented in the
collections. There are 39 specimens, of which the following
were measured: BMNH EE4211, EE4250, EE4277,
EE4279-80, EE4283, EE4285, EE4287, EE4291,
EE4898-99, EE4301, EE4304-06, EE4308, EE4310.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

OCCURRENCE. This species is found in the western Oman

Mountains at the following levels:

Jebel Huwayyah, section 1: beds 3-5 (1).

Jebel Buhays, section 1: loose, derived from lowest few
metres of section (19).

Jebel Thanais: lowest few metres of the Simsima Formation
(11).

Jebel Rawdah, section 2: bed 14 (1); bed 15 (2); bed 21 (3);
loose, mid-section (1).

Jebel Rawdah, section 3: loose (4).

Jebel Rawdah, section 4: beds 8/9 (1); bed 13 (1); loose in
scree (1).

_ Outside Oman, the species is known from Libya, Saudi
Arabia, southern Iran and Assam, India.

DIAGNOsIS. A species of Conulus with a rounded to strongly
fusiform peristome which is not sunken. Periproct situated
relatively low on the posterior surface, not visible from
above. Pore-pairs adorally arranged triserially. Apical disc
with genital plates 3 and 4 always in contact. Test profile
subconical.

DESCRIPTION. Tests range from 17 to 45 mm in length and
are ovoid to rounded pentagonal in outline. Test width is
| 82-96% of test length (mean = 88%, SD = 3:2%, N = 18)
_ and the widest point coincides with the posterior part of the
_ antero-lateral ambulacra. Test height is 64-81% of test length
| (mean = 72%, SD = 4.9%, N = 20) and in profile the test
has a broad, flat base and is rounded subconical above (PI.
20, figs 11-16).
The apical disc lies centrally and is tetrabasal. Genital plate
| 2abuts the other three genital plates but never reaches ocular
' plate 4 to separate genital plates 3 and 4 (Figs 47C-E).
Genital plates 4 and 1 are always in contact behind the
' madreporite. There appears to be some degree of differentia-
tion in gonopore size, suggestive of sexual dimorphism.
Ambulacra are straight and compound in the pyrinoid
| style. Above the ambitus pore-pairs are strictly uniserial, but
/ below the ambitus they become offset into three discrete
columns and these continue to the peristome edge (PI. 19, fig.
/ 9; Pl. 20, fig. 11; Fig. 48B).
The periproct lies on the posterior border, close to the base
(Pl. 20, fig. 15). It is tear-drop shaped, being pointed
adapically. Its height is 28-40% of the test height (mean =
35%, SD = 3:2%, N = 20) and its width is 55-77% of its
height (mean = 65%, SD = 6:2%, N = 19). The base of the
periproct lies 3-17% of the test height above the base (mean
= 11%, SD = 3-6%, N = 20). The periproct is just visible
from beneath, but is not seen from above (PI. 19, figs 8, 9).
The peristome is oval to fusiform in outline and is not
\invaginated, although the oral surface may be slightly

‘depressed towards the peristome. Its length is 14-23% of the
“test length (mean = 20%, SD = 2:7%, N = 13). Its width is
56-81% of its length (mean = 70%, SD = 8-1%, N = 13). It
is oblique, with its long axis orientated from interambulacrum
3 to ambulacrum I (PI. 19, fig. 9).

|REMARKS. The distinction between this species and ‘Globa-
tor’ bleicheri is detailed above. This species comes closest to
— Conulus giganteus Noetling (C. gigas Cotteau, C. ataxaensis

Cotteau) but in this species complex the peristome is less
ovoid and is distinctly more invaginated than C. gigas at least.
Whether the Middle Eastern species turn out to be suffi-
ciently distinct remains to be seen.

189

Order CASSIDULOIDA Claus, 1880
Family CLYPEOLAMPADIDAE Kier, 1962

DIAGNOsIs (following Kier, 1962). Cassiduloids with a
domed test and flat base. Petals long and straight, apical disc
tetrabasal, periproct inframarginal and transverse, bourrelets
with three or more columns of pores and with buccal pores
present.

TYPE GENUS. Clypeolampas, type ovatus

Lamarck, 1816.

Goeoes (C-

OTHER GENERA INCLUDED. Hungaresia Szorenyi, type spe-
cies Hungaresia hungarica Szorenyi (junior synonym of Cly-
peolampas ovum Grateloup); Vologesia Cotteau & Gauthier,
type species V. tataosi Cotteau & Gauthier.

REMARKS. Lambert (1919) claimed that the original figure
and description of Clypeolampas ovatus Lamarck (1816: 22)
was inadequate for certain identification, and thus used the
name Clypeolampas leskei Goldfuss (1829) as the oldest
available name. However, Kier (1962: 190) accepted C.
ovatus Lamarck as a valid designation and sunk C. leskei
Goldfuss in synonymy.

There are eighteen nominal species assigned to the three
genera listed above. The differentiation of the three genera
is, however, unclear. Vologesia was established by Cotteau &
Gauthier (1895: 65) for a late Cretaceous species from Aftab,
Iran, V. tataosi Cotteau & Gauthier. This is based on a single
small individual, subcircular in plan view, with poorly devel-
oped bourrelets and the peristome positioned close to the
anterior border. No other species were included in this genus
until Lambert (1919) revised the then known members of
Clypeolampas. He separated species into two groups; those
with uniform aboral tuberculation, all tubercles being scro-
biculate, and those forms which had a second kind of aboral
tuberculation composed of nonscrobiculate pustules. The
former he assigned to Vologesia, the latter to Clypeolampas.
Lambert (1919) placed the following species in Vologesia: C.
ovum Grateloup, C. acuta Desmoulins, C. conica Arnaud, C.
toucasit Lambert, C. gossauviensis Lambert and a small
undescribed Maastrichtian form. In Clypeolampas Lambert
placed C. leskei Goldfuss [=C. ovatus Lamarck], C. perova-
lis, Arnaud, C. orbicularis Arnaud, C. lestelei Cotteau, C.
vishnu, Noetling, C. douvillei Lambert and C. mengaudi
Lambert.

In 1955, Szorenyi erected the genus Hungaresia for the new
Santonian species H. hungarica Szorenyi of Hungary. This
species appears to be identical in many important respects to
the common Santonian species Clypeolampas ovum Grate-
loup, and Kier (1962: 191) synonymized the two genera,
making Hungaresia a junior synonym of Vologesia.

Kier (1962, 1966) followed Lambert’s generic differentia-
tion, distinguishing Clypeolampas from Vologesia by its non-
scrobiculate pustules developed adapically, and by its better
developed floscelle. However, Kier’s concept of Vologesia
was based not on the type species but on Clypeolampas ovum
Grateloup and is thus misleading.

There are very few stable characters on which to subdivide
the group, the following being amongst the most informative.

(1) Aboral pustules developed. In some species there are
characteristic pustules over the adapical surface that are
slightly larger than the normal tuberculation and give the
surface a rugose appearance. These are not tubercles for
spine articulation, as they have no articular surface or sur-

190

rounding scrobicule. Instead they resemble the pustular cal-

cite formed in species of Conulus or Echinoneus (Smith

1980). Their function is unknown but it may be to do with

deterring parasitic and commensal settlement. Pustules are

developed in a number of species, including the type C.

ovatus Lamarck.

(2) The position of the peristome seems highly stable and
distinctive. In Vologesia tataosi and V. rawdahensis the
peristome lies close to the anterior border, with the anterior
edge lying between 20 and 25% of the test length from the
anterior. In almost all other species the peristome is subcen-
tral, lying between 30 and 40% of the test length from the
anterior. Only one species, Clypeolampas toucasi Lambert is
intermediate in this respect, with its peristome between 25
and 30% of test length from the anterior.

(3) Elongation of the peristome. Only Vologesia rawdahen-
sis Ali has such a laterally elongate peristome. In other
species the peristome is suboval.

(4) The degree to which the floscelle projects as prongs
over the peristome is to some extent size dependent, with
more prominent floscelles in larger individuals. However, at
a similar size it is apparent that Vologesia tataosi, V. rawda-
hensis and Clypeolampas toucasi show virtually no floscelle
development, whereas C. ovatus and its synonyms and C.
perovalis have very pronounced floscelle projection. In C.
lestelei, C. ovum, and C. conicus as well as probably C.
helios, the floscelle is slightly swollen but not projecting.

(5) The periproct is usually unambiguously positioned on
the flat oral surface, but in a few species such as C. helios and
C. ovum, the periproct lies subambitally because of the
strongly inflated test profile.

(6) The apical disc is definitely tetrabasal in C. /estelei, C.
ovum and Vologesia rawdahensis, but monobasal in C. ova-
tus.

(7) The arrangement of pores in the bourrelets, though to
some extent size dependent, offers some differentiation. In
C. ovatus there are many pores irregularly scattered between
inner and outer columns in each half ambulacrum. By con-
trast C, ovum and Hungaresia hungarica have only the inner
and outer series of pores. If C. conicus is just a variety of C.
ovum as is suspected, then some specimens may have a few
pores forming a mid row. The same is true of Vologesia
rawdahensis where individuals have either two or three rows.

From this the following supraspecific taxonomy is pro-
posed:

Clypeolampas Lamarck: Large forms with moderate to well
developed bourrelets, subcentral peristome and aboral
calcite pustules. Species included; C. ovatus Lamarck
(includes C. leskei Goldfuss, C. mengaudi Lambert, C.
douvillei Lambert, C. orbicularis Arnaud), Upper
Campanian-Maastrichtian of Spain, southern France, Tur-
key; C. perovalis Arnaud, Lower and Middle Campanian
of Gironde, France; ?C. lestelei Cotteau, ?Danian of Saint
Cirac, Ariége, France.

Vologesia Cotteau and Gauthier. Distinguished from Cly-
peolampas by its anterior peristome without floscelle devel-
opment. Type species V. fataosi Cotteau & Gauthier,
Upper Senonian of Louristan, Iran; V. rawdahensis Ali,
Maastrichtian of the Oman Mountains; V. toucasi (Lam-
bert), Campanian of the Pyrenees, France.

Hungaresia Szorenyi. Smaller, ovoid forms with subcentral,
pentagonal peristome with swollen but not projecting flos-
celles, no aboral pustules and subambital rather than fully
adoral periproct. Type species, H. ovum (Grateloup)

A.B. SMITH

[includes H. hungarica Szorenyi], Upper Santonian of
France, Pyrenees, Hungary. Other species included: H.
helios (Noetling), ?Maastrichtian, Mari Hills, Baluchistan.

Unplaced taxa: Clypeolampas conicus Arnaud (U. Santonian|_
to L. Campanian, SW France) and C. acuta Desmoulins
(pores in bourrelets shown as forming two columns only).
material of these species has not been seen and so they
cannot be placed with any confidence from the published!
descriptions and figures.

C. vishnu Noetling: based on a single worn specimen, inad-
equately known nomen dubium.

C. gossaviensis Lambert: based on a single, poorly preserved
specimen and indeterminate from description and figure}
nomen dubium.

Genus VOLOGESIA Cotteau & Gauthier, 1895

TYPE SPECIES. Vologesia tataosi Cotteau & Gauthier, 1895, i
by original designation.

OCCURRENCE. Late Cretaceous (‘Upper Senonian’) of Iran;
Maastrichtian of the United Arab Emirates and Oman.

DIAGNOSIS. Clypeolampadids with an anteriorly positioned!
peristome lying 20-30% from the anterior border. Peristome
wide, pentagonal with bourrelets hardly developed. Phyl-
lodes with two or three rows of pores in each half ambu
lacrum. Apical disc tetrabasal.

REMARKS. Vologesia is distinguished from Clypeolampas b
its lack of aboral pustules and its poorly developed floscelles _
and bourrelets. It is distinguished from Hungaresia by its lac
of floscelles, its more transverse peristome and its mor
anterior peristome.

Pl. 21, figs 1-5; Figs

Vologesia rawdahensis Ali, 1989 :
49-5].

1989 Vologesia rawdahensis Ali: 406, fig. 5 (13).

TyPEs. Three specimens in the Geology Museum, United
Arab Emirates University, Al Ain, United Arab Emirates.

MATERIAL STUDIED. Seven specimens, of which fiv
(BMNH EE3383-85, EE4326, EE4329) are well enoug
preserved to be included in the biometric analysis.

OCCURRENCE. This species is known only from the Simsim

Formation of the western margins of the Oman Mountains.

Specimens were found at the following localities and hori

zons:

Jebel Buhays, section 1: loose in the scree, derived from th
lowest few metres of Simsima Formation (5).

Jebel Buhays, section 2: loose in the scree, derived from th
lowest 2 m of Simsima Formation (1).

Jebel Thanais: lowest 2 m of Simsima Formation (1).

Jebel Rawdah, section 2: bed 19 (1).

Jebel Rawdah, section 3: bed 8 (1).

DESCRIPTION. Tests are flat-based and rounded to subconi
cal in profile (Pl. 21, figs 1-4), with a relatively sharp ambitu
situated low down. In outline the test is ovoid with a rounde
anterior and a distinctly more pointed posterior. Tests rang
in length from 46 to 68 mm. Test width is 75-77% of tes
length and test height 53-62% of test length (Fig. 49). The
tallest part of the test is central or slightly anterior of centre.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

@ Test width (mm)
@ Test height (mm)

50 60 70
Test length (mm)
30
@ to apical disc
@ to peristome

Distance from anterior (mm)
N
°

=
v2
°o

60 70
Test diameter (mm)

Length of petal (mm)

60
Test length (mm)

Fig. 49 Biometric data for Vologesia rawdahensis Ali.

The apical disc is tetrabasal (Fig. 51B), with the
madreporite very large and occupying the centre. Other
genital plates are much smaller and pushed far out into the
adjacent interambulacra. The apical disc lies 37-39% of test
length from the anterior border (Fig. 49).

Petals are relatively long and straight or only very slightly
powed. They are open distally and extend most of the
listance towards the ambitus. The posterior petals are always
blightly longer than the anterior three petals. They are
pomposed of an inner circular pore and a highly elongate
buter slit-like pore, joined by a furrow. There are 51 pores in
1 petal column in a 53 mm individual and 57 in a 68 mm
individual. Pores below the petal are all single.

The peristome is pentagonal in outline and very much
vider than long (width is 1-7-2-0 times greater than length).

191

It is straight sided and only slightly invaginated (PI. 21, fig. 5;
Fig. 50) and bourrelets are hardly developed. It lies 21-27%
of the test length from the anterior border. Phyllodes are
relatively short and are not depressed in the slightest. In some
individuals there are only two columns of pores in each half
ambulacrum, whereas in others there are three columns. The
outer series is composed of about 12 pores, the inner series of
7 or 8 pores and the mid series, where present, of 4 or 5 pores
(Fig. 51). The inner and middle series of pores are borne on
occluded plates. Buccal pores are present.

The periproct lies on the oral surface at the posterior. It is
oval in outline, approximately twice as wide as long and
approximately the same size as the peristome.

Aboral tuberculation is fine and uniform, oral tubercula-
tion slightly coarser and becoming less dense towards the
midline. There is a broad tubercle-free band down the
midline in the posterior interambulacrum running between
the peristome and periproct. This is lightly pitted (PI. 21, fig.
5).

REMARKS. This species was erected by Ali (1989) on the
basis of three specimens from Jebel Rawdah, Oman. It differs
from V. tataosi Cotteau & Gauthier, from a similar horizon in
southern Iran, by being more elongate and pointed posteri-
orly, and by having a smaller, more transversely elongate
mouth (if the original figures of this species are true to life).
The Iran species was, however, based on a single individual
27 mm in test length, and there is the possibility that with
more material the two species may eventually prove to be
synonymous.

Family FAUJASIIDAE Lambert, 1905
Genus FAUJASIA d’Orbigny, 1856

TYPE SPECIES. Pygurus apicalis Desor, by subsequent desig-
nation of Lambert & Thiery, 1921: 273.

OTHER SPECIES INCLUDED. Only one other species, Faujasia
eccentripora Lees. Two species previously ascribed to Fauja-
sia were transfered to other genera by Kier (1962): F. faujasi
(Desmoulins) to Eurypetaium and F. chelonium Cooke to
Domechinus.

DIAGNOSIS. Small ovoid cassiduloids with a monobasal api-
cal disc in which the genital pores open through interambu-
lacral plates. Petals broad, closed distally and strongly
petaloid in form. Periproct small, circular and inframarginal.
Peristome small, anterior with short, arcuate phyllodes in two
columns and with buccal pores, and strongly projecting
bourrelets.

OCCURRENCE. Maastrichtian of Belgium, France, Oman and
the United Arab Emirates.

REMARKS. Kier (1962: 137), in discussing this genus, was
uncertain whether F. eccentripora Lees truly belonged here,
since its apical disc plating had never been described. As
shown below, this species has an identical arrangement of
gonopores opening within the adapical portion of the inter-
ambulacra as characterizes F. apicalis Desor and thus is
clearly closely related. The derived position of gonopores
outside apical disc plating distinguishes these two species
from all other cassiduloids.

|
|
192 A.B. SMITH |

“f o

oo "<<?
: tC One:
OOS G Cre,

wf OD
CAGES SSogu

PLATE 21

Figs 1-5 Vologesia rawdahensis Ali. 1-3, 5, BMNH EE4326; 1, oral, x 1; 2, apical, x 1; 3, lateral, x 1; 5, detail of peristomial region, X
2-5. Jebel Thanais, lowest 2 m of the Simsima Formation. 4, BMNH EE4329, lateral, x 1. Jebel Buhays, section 1; loose in the scree |
derived from the lowest 3 m of the Simsima Formation.

Figs 6-13 Zuffardia morgani (Cotteau & Gauthier). 6-8, BMNH EE3789; 6, oral; 7, lateral; 8, posterior, all x 2. Jebel Rawdah, section 2, |
bed 11. 9, BMNH EE3791; apical, x 2. Jebel Rawdah, section 2, bed 21. 10-13, Topotype specimen in the Morgan Collection, Museum
d’Histoire Naturelle, Paris; 10, apical; 11, oral; 12, posterior; 13, lateral; all x 1-5. Senonian, Dah-e-Rouh Davl, Iran. |

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

Fig. 50 Camera lucida drawings of Vologesia rawdah
bars = 5 mm.

Ali. A, oral plating, BMNH EE3384; B, adapical plating, BMNH EE4326. Scale

ay
a) Nd ,
ct a

Fig. 51 Camera lucida drawings of Vologesia rawdahensis Ali. A, peristome and surrounding phyllodes, BMNH EE3384; B, apical disc
plating, BMNH EE4326. Scale bar 1 mm.

193

194
40 Ss
@ Test width r
@ Test height a
E
E
Test length (mm)
15
@ Lenath
@ Width Ss
£
£
~~
w
~
@
a
_
gS
o
Cd
c
<

Test length (mm)

Fig. 52 Biometric data for Faujasia eccentripora Lees.

Faujasia eccentripora Lees, 1928 Pl. 22, figs 5—14; Figs
52-55

1928 Faujasia eccentripora Lees: 661, pl. 46, fig. 2.
1989 Faujasia eccentripora Lees; Ali: 403, fig. 4 (25).

Types. Holotype, BMNH_ E18347, paratypes BMNH
E1834446, E18348, from the late Cretaceous (?Maastrich-
tian) of Jebel el Malih, Oman.

MATERIAL STUDIED. 465 specimens were collected for study,
of which 29 (BMNH EE3823, EE3825—26, EE3832-34,
EE3836-39, EE3851, EE3855-61, EE3866-67, EE3872-76,
EE3878-79, EE3882, EE3897) were measured for the bio-
metric analysis given here.

OCCURRENCE. This species is known only from Oman and

the United Arab Emirates. It occurs abundantly at the

Loftusia levels at Jebel Huwayyah and in the lowest 8 m of

calcarenitic limestones at Jebel Rawdah 2. Elsewhere it is

rare. It was collected at the following localities and strati-
graphic levels:

Jebel Aqabah: bed 1 (6, all juvenile).

Jebel Huwayyah, section 1: bed 9 (6); beds 10-11 (33).

Jebel Huwayyah, section 2: bed 3 (1).

Jebel Rawdah, section 1: top of bed 2 (5); top of bed 4 (2);
loose in scree at base (6).

Jebel Rawdah, section 2: bed 4 (14); bed 5 (2); beds 6-8 (92);
bed 10 (1); bed 11 (101); bed 14 (7); bed 16 (3); bed 19
(19); bed 20 (2); bed 21 (22); bed 22 (3); beds 23-25 (1);
bed 25 (3); loose, from lower part of succession (beds 2-11)
(112); loose in scree from higher part of succession (13).

A.B. SMITH

® anterior to peristome %
@ Anterior to apical disc

Distance (mm)

10) 10 20 30 40
Test length (mm)

& Peristome
@ Periproct

Length (mm)

te) 10 20 30 40
Test length (mm)

Jebel Rawdah, section 3b: bed 2 (3); bed 8 (1); bed 9 (3).
Jebel Rawdah, section 4: bed 2 (1); bed 4 (2); beds 21-22 (1).

DESCRIPTION. Tests are shield-shaped in outline with a
rounded anterior, a projecting pointed posterior and two
marked angles coinciding with the posterior ambulacra (PI.
22, figs 5-14; Fig. 53). Tests range in length from 8 to 37-5
mm. They have a rounded ambitus and slightly convex oral
surface, with a marked sternal keel. The test is widest
posterior of midlength and maximum width is 92-106% test
length. The upper surface is low and rounded, never peaked.
Test height is 50-77% of test length, being greatest in small
individuals and progressively decreasing through growth (Fig.
52).

The apical disc lies 30-44% of test length from the anterior
border (mean = 38%, N = 29). It is more central in larger
forms. The disc is monobasal with the entire apical region
occupied by the madreporite. There are four gonopores but
these lie outside the apical disc, opening instead within the
interambulacra and separated from the madreporite by one
or two interambulacral plates (Fig. 54).

The petals are strongly inflated and closed distally, being
widest at midlength (PI. 22, figs 6, 10, 13; Fig. 53B). All five
are similar in size and the anterior three extend some 70-80%
of the way from the apex to the ambitus. Both pores in the
pore-pair are circular or subcircular and united by a furrow.
Pores are all single beneath the petals.

The peristome is small, only about 10% of the test length in
diameter. It is rounded quinquestellate in outline and
approximately as wide as broad. It lies 20-30% of the test
length from the anterior border (mean = 25%, N = 29). The

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 195

PLATE 22

Figs 1-4 Zuffardia morgani (Cotteau & Gauthier). BMNH EE4325; 1, apical; 2, oral; 3, lateral; 4, posterior; all x 2. Jebel Rawdah, section
1, bed 3.

Figs 5-14 Faujasia eccentripora Lees. 5-7, BMNH EE3821; 5, oral; 6, apical; 7, posterior; all x 2. Jebel Rawdah, section 2, bed 14. 8-11,
BMNH EE3826; 8, lateral; 9, oral; 10, apical; 11, posterior; all x 2. Jebel Rawdah, section 2, loose in scree at level of bed 14. 12-14,
BMNH EE3825; 12, oral; 13, apical; 14, lateral; all x 2. Jebel Rawdah, section 2, bed 11.

196

Fig. 53 Camera lucida drawings of Faujasia eccentripora Lees. A, BMNH EE3882, oral surface; B, BMNH EE3824, adapical surface. Scale

bar = 5 mm.

basicoronal plates of the interambulacra are rather narrow
and elongate (Fig. 55), as are the next two pairs of plates in
the posterior three interambulacra (Fig. 54). The bourrelets
are rather square-sided and blunt-ended and project strongly
(Pl. 22, figs 5, 9, 12). The phyllodes are short and broad and
are sunken. Pores are strongly arcuate with 8 to 12 pores in
the outer series and only 2 or 3 in the inner series (Fig. 55).
The inner series of pores are well separated from the peris-
tome, but there are two pairs of sphaeridial pits adorally (Fig.
55). Miniscule buccal pores are present but are much smaller
than other pores in the phyllodes and are clearly rudimentary
only.

The periproct is small and circular, width being 85—122% of
its length. It lies close to the posterior margin on the oral
surface at the end of the sternal ridge and faces slightly
posteriorly (Pl. 22, figs 7, 11).

Tuberculation is fine and uniform aborally and slightly
coarser adorally. There is a narrow naked zone that extends
from just posterior of the posterior bourrelet half way
towards the periproct.

REMARKS. This species was first erected by Lees (1928) for
specimens from Jebel el Malih, Oman. It differs from Fauja-
sia apicalis Desor, from the Maastrichtian of Belgium, the
Netherlands and France in a number of important respects.
F. apicalis has shorter petals, a sharper ambitus and is more
pointed apically in profile. Furthermore, its peristome is
much less anterior and the oral surface is flatter and lacks
such a pronounced sternal keel.

Genus ZUFFARDIA Checchia-Rispoli, 1917

TYPE SPECIES. Pseudocatopygus sanfilippoi Checchia-
Rispoli, 1914, by original designation.

OTHER SPECIES INCLUDED. In addition to the type species,
there are four nominal species, three of which have previ-

A.B. SMITH

==)

=,
7
=,

Vea

ZS
BZ
Za

/ we
ae

Fig. 54 Camera lucida drawings of apical disc plating of Faujasia
eccentripora Lees. A, BMNH EE3824; B, BMNH EE3826. Scale
bar = 1 mm.

Fig. 55 Camera lucida drawing of phyllode plating in Faujasia
eccentripora Lees, BMNH EE3902. Scale bar = 1 mm.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

@ Test width
@ Test height

Distance (mm)

10 20 30 40
Test length (mm)
10
® Distance from base to periproct
®@ Periproct height
8 mi
te. o °
-
e
E 6 + &*,
~

2
Test height (mm)

Fig. 56 Biometric data for Zuffardia morgani (Cotteau & Gauthier).

ously been assigned to this genus; Catopygus rohlfsi Krum-
beck, 1906, Catopygus boucarti Lambert & Thiéry, 1925,
Catopygus morgani Peron & Gauthier, 1895, Zuffardia
creullii Checchia-Rispoli, 1933. All are here treated as syn-
onymous with C. morgani, the oldest available name.

DIAGNOsIs. Oval test with a monobasal apical disc, broad
petals of equal length, a posterior longitudinal periproct and
a longitudinal peristome with short phyllodes and buccal
pores.

OCCURRENCE. The genus occurs in the late Cretaceous of
North Africa and the Middle East. It is recorded from the
Maastrichtian of Libya, Algeria and Oman and the United
Arab Emirates, and from the upper Senonian (?Maastrich-
tian) of southern Iran.

_ REMARKS. This genus resembles Catopygus Agassiz, 1836, in

its general shape and form but differs from that genus in
having a monobasal apical disc and single pores below the

_ petals. It also appears rather similar to the Tertiary genus

Kephrenia Fourtau, 1909, having very similar phyllodes and

| longitudinal peristome. It differs from Kephrenia in periproct

shape, Kephrenia having a small transverse periproct as

| opposed to the longitudinal periproct of Zuffardia. It also
| shares many characteristics in common with Faujasia, includ-

ing its phyllode and bourrelet structure, small periproct

, positioned low on the posterior face and apical disc plating. It

differs from Faujasia in having gonopores confined to genital
plates, and in being much more oval in shape.

Only two species are recognized here, the type Z. sanfilip-
poi and Z. morgani (Cotteau & Gauthier).

Peristome length (mm)

197

gs Anterior to peristome
@ Anterior to apical disc

ona

20 30

Test length (mm)

20
Test length (mm)

30 40

Zuffardia morgani (Cotteau & Gauthier, 1895) PI. 21,
figs 6-13; Pl. 22, figs 1-4; Figs 56-59

1895 Pseudocatopygus Morgani Cotteau & Gauthier: 60,
pl. 9, figs 6-9.
21906 Catopygus Rohlfsi Krumbeck: 87, pl. 7, fig. 4.
1914 Pseudocatopygus rohlfsi Krumbeck; Checchia-
Rispoli: 301, pl. 1, fig. 3.
1925 Catopygus boucarti Lambert; Lambert & Thiery:
587, pl. 13, figs 6-9.
1933 Zuffardia creullii Checchia-Rispoli: 4, pl. 1, figs 1-4.
1939 Catopygus boucarti Airaghi: 258.
1981 Zuffardia rohlfsi boucarti (Lambert); Roman, in
Amard et al.: 112.
1987 Zuffardia aff. rohlfsi boucarti (Lambert); Zhagbib-
Turki: 257.
1989 Zuffardia sanfilippoi Ali: 406, fig. 5 (67).
Types. The holotype is the single specimen figured and

described by Cotteau & Gauthier (1895). It has not been
located. However, another specimen from the same locality
and in the Morgan collection is figured here (Pl. 21, figs
10-13).

MATERIAL STUDIED. Seventy one specimens were collected,
of which 28 (BMNH EE3382-85, EE3531, EE3789-93,
EE3796-804, EE3806, EE3808, EE3812-17, EE4325,
EES050) were used for the following biometric analysis.

OCCURRENCE. The species occurs In Tunisia, Algeria and
Libya as well as in southern Iran and the United Arab
Emirates and Oman. It is apparently restricted to the Maas-

198

NE
RNY Oa 4
NY Lee

Fig. 57 Camera lucida drawing of apical disc plating in Zuffardia
morgani (Cotteau & Gauthier), BMNH EE3791. Scale bar = 1
mm.

trichtian. In the study area described here, the species is

found only at Jebel Rawdah, at the following localities:

Jebel Rawdah, section 1: bed 3 (5).

Jebel Rawdah, section 2: bed 11 (47); bed 19 (5); bed 21 (5);
bed 25 (1); loose, mostly derived from beds 2-11 (48).

Jebel Rawdah, section 4: bed 2 (1).

DIAGNOsIS. A species of Zuffardia with a rather flat base
and a periproct that becomes progressively more adoral as
test size increases. Test shape highly variable, either inflated,
keeled or depressed.

DESCRIPTION. Tests ovoid in shape ranging from 10-8 to 33-7
mm in length. In plan view they are rounded anteriorly and
slightly pointed posteriorly (Pl. 21, fig. 9). The ambitus is

A.B. SMITH

rounded and in profile the base is flattish and the upper
surface ranges from low-domal or even flat, to conical and
pointed at the apex (PI. 21, fig. 7; Pl. 22, fig. 3). Test width is
83-97% of the length (mean = 90%, N = 25) and test height
55-91% of the length (mean = 71%, smaller individuals
being proportionately taller, Fig. 57).

The apical disc is monobasal and positioned 33-47% of the
test length from the anterior border (mean = 40%, N = 24;
Fig. 56). The madreporite is large and its pores extend to, and
partially enclose, the four gonopores (Fig. 57). There appears
to be a distinct sexual dimorphism in gonopore size, with
large individuals having either large, closely spaced gonop-
ores some 0-3-—0-5 mm in diameter (female?) and others of
comparable size having only small (0-1 mm diameter) gonop-
ores (male?).

Petals are lanceolate and converge distally but do not close
(Pl. 22, fig. 1; Fig. 58B). Petals Il, II and IV are about
70-80% the length of the posterior petals. The two pores in
each pair are approximately circular and connected by a
groove. The three anterior petals extend approximately 70%
of the distance to the ambitus, the posterior pair only some
60% or so. In middle to large individuals the interporal zone
is 1-0-1-5 times as broad as an individual pore zone. Pores
below the petals are all single.

The peristome is pentagonal in outline and slightly longer
than wide (length is 1-2—-1-5 times its width; mean = 1-3). It
lies 33-38% of test length from the anterior border. Phyllodes
are short and arcuate, with two series of pores in each column
(Fig. 59). There are 6 or 7 pores in the outer series and only 1
or 2 in the inner series. There are also two pairs of sphaeridial
pits in each half ambulacrum situated perradially. A small
number of occluded plates are developed and buccal pores
are present. The phyllodes are not depressed but remain flush
with the test. The bourrelets project outwards strongly but do
not indent the peristome (PI. 21, fig. 6; Pl. 22, fig. 2).

|
i

Soon

PRS
1]
Sas
ee
S3s=
aS
SS.
Za =)
ST ae

int

Fig. 58 Camera lucida drawings of plating in Zuffardia morgani (Cotteau & Gauthier). A, BMNH EE3385, oral plating; B, BMNH EE3384,

adapical plating. Scale bar = 2 mm.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

The periproct lies on the posterior face, relatively high in
juveniles but becoming progressively lower in larger speci-
mens (Fig. 56). The base of the periproct lies 50% of test
height above the base in small individuals, but can lie as low
as 22% of test height in the largest individuals. There is,
however, considerable variation in the height of the periproct
within the sample. The periproct is relatively small and taller
than wide (height = 1-3-2:5 times greater than width).
Typically, there is a slight adapical projection and rim devel-
oped around the periproct.

Tuberculation is fine and uniform aborally and slightly
coarser adorally. There is a narrow sternal naked zone on the
oral surface behind the peristome.

REMARKS. This species was first described from the ‘Upper
Senonian’ (probably Maastrichtian) of Derre-i-Chahr,
southern Iran, by Cotteau & Gauthier (1895: 60) under the
name Catopygus morgani. Cotteau & Gauthier distin-
guished it from other similar species by its inflated shape
and convex oral surface. Although their figure indicates a
peristome that is as broad as long, in all other respects it
falls exactly within the range of the United Arab Emirates
population described here (Fig. 57). Should the Iranian
population prove to have a consistently more equant
peristome, then it clearly should be treated as a distinct
species. However, the figures cannot be considered
entirely reliable and this difference is not apparent in
topotype material (PI. 21, figs 10-13). Consequently, for
the present the two forms are synonymized.

In 1906 Krumbeck erected the species Catopygus rohlfsi on
the basis of a specimen from the late Cretaceous of Libya.
Closely comparable material from Algeria was given the
name Catopygus boucarti by Lambert & Thiéry (1925). Based
on a large collection of specimens from the Maastrichtian of
Libya, Airaghi (1939) was later able to synonymize Lambert’s
Catopygus boucarti and Krumbeck’s Catopygus rohlfsi. The
large populations of Zuffardia boucarti Lambert, described
from Algeria by Roman (in Amard ef al. 1981) also fall within

_ the range of the material described here, though they do not
_ achieve a size greater than about 2 cm (Libyan specimens

cover an identical range to the population described here).

Fig. 59 Camera lucida drawing of adoral plating of Zuffardia
morgani (Cotteau & Gauthier), BMNH EE4325. Scale bar = 1
mm.

199

Thus there can be little doubt that these populations are all
synonymous. However, there remains slight doubt over
whether Krumbeck’s original specimen is conspecific. Roman
(in Amard ef al. 1981) pointed out that Krumbeck’s (1906)
original figure shows a specimen whose apical disc lies well
displaced towards the anterior and whose periproct is shown
as being visible from above. I suspect that this is simply due to
the Krumbeck’s specimen being slightly crushed and dis-
torted, or possibly simply being illustrated from an oblique
angle. But the type specimen has not been located and thus
the validity of this species must remain uncertain.

In 1914 Checchia-Rispoli described a more spherical form
with a strongly inflated oral surface under the name
Pseudocatopygus sanfilippoi. There are three specimens, the
largest of which is 42 mm in length. This form has a much
higher periproct than is found in my populations (Fig. 57),
and corresponds to the juvenile positioning (as does the more
spherical test shape). This lies well outside the range of form
found in Z. morgani from either the Algerian or the United
Arab Emirates populations, and clearly represents a distinct
species. Checchia-Rispoli (1917) made this the type species of
his new genus Zuffardia.

The only other species assigned to Zuffardia is Z. creullii
Checcia-Rispoli (1933). This is based on a single individual 33
mm in length that corresponds to a more inflated variant of
Z. morgani. Biometrically it falls more or less within the
range of variation encountered in the population described
here. It too, therefore, is synonymized with Z. morgani.

Unnamed Family
Genus PYGUROSTOMA Cotteau & Gauthier, 1895

TYPE SPECIES. Pygurostoma morgani Cotteau & Gauthier,
1895, by original designation.

OTHER SPECIES INCLUDED. None.

OCCURRENCE. Late Cretaceous of Iran and the Oman
Mountains.

DIAGNOSIS. Large, oval and rather flattened cassiduloid with
a narrow, elongate and invaginated inframarginal periproct.
Peristome quinquestellate and subcentral with strong phyl-
lodes containing many occluded plates and pores. Buccal
pores present, well separated from the peristome. Apical disc
tetrabasal and subcentral. Petals long and well developed,
converging distally. Naked zone on the oral area between the
peristome and periproct, and also extending anterior of the
peristome in ambulacrum III.

REMARKS. Kier (1962) included Pygurostoma in the family
Faujastidae on account of its phyllode structure. However, as
Kier pointed out, this genus is rather different from other
Faujasiidae. In particular the appearance of the periproct,
which is strongly rostrate adapically and invaginated adorally,
indicates that it is probably derived from a form with a
posterior elongate periproct and weak anal canal. This is very
different from the periproct seen in Faujasia and most other
faujasiids. In test shape, peristome position and size, floscelle
development, petal form and the distribution of naked zones,
Pygurostoma undoubtedly comes closest to Parapygus
Pomel, 1883. Parapygus has a narrow, vertically elongate
periproct that lies posteriorly and in large forms is just visible
adorally. Although many species have just two columns of
pores in each half ambulacrum of the phyllodes, the type

200

species, P. cotteauanus (d’Orbigny) has a median zone of
pores also. Thus, in many respects Pygurostoma is no more
than an hypertrophic form of Parapygus in which the phyl-
lodes have increased numbers of median pores and the
periproct is inframarginal rather than marginal.

Only one other species was included in this genus by Kier
(1962), Pygurostoma pasionensis Cooke, 1949, from the late
Cretaceous of Guatemala. However, this species has its
periproct flush with the test and not invaginated. I suspect
this is not cogeneric.

Pygurostoma morgani Cotteau & Gauthier, 1895 PI. 23,
figs 1-4, 7; Figs 60-63

1895 Pygurostoma morgani Cotteau & Gauthier: 52, pl. 8,
figs 1-5.

1962 Pygurostoma morgani Cotteau & Gauthier; Kier: 135,
pl. 19, figs 1-3, text-fig. 117.

1992 Pygurostoma morgani Cotteau & Gauthier, Ali: 72,
fig. 4.

Types. The whereabouts of Cotteau & Gauthier’s specimens
is unknown. A topotype specimen from the Morgan collec-
tion is illustrated (Pl. 23, fig. 1).

MATERIAL STUDIED. Forty specimens, of which 15 (BMNH
EE3291-92, EE3296, EE3302-10, EE3315, EE3317,
EE3322) were measured.

OCCURRENCE. The species was first described from the
‘Upper Senonian’ of Aftab, Louristan, southern Iran. In this
study specimens were most common in the lowest few metres

@ Peristome
@ Apical disc

Diameter (mm)

30 40 50 60

Test diameter (mm)

Test height (mm)

30 40 50 60

Test diameter (mm)

Fig. 60 Biometric data for Pygurostoma morgani Cotteau & Gauthier.

Width at ambitus (mm)

(mm)

A.B. SMITH

of the Simsima limestones. Fifty specimens were collected or

noted at the following localities and horizons:

Jebel Buhays, section 1: loose in the scree, derived from the
lowest few metres of Simsima Formation (26).

Jebel Buhays, section 2: loose in scree, derived from lowest 2
m of the Simsima Formation (2).

Jebel Thanais: lowest 2 m of Simsima Formation (1).

Jebel Huwayyah, section 1: beds 14/15 (3).

Jebel Rawdah, section 2: bed 14 (3); bed 18 (1); bed 19 (2);
bed 21 (7); bed 26 (2); loose in scree (1).

Jebel Rawdah, section 4: bed 19 (1).

Jebel Faiyah, section 1 (southern tip): bed 8 (1).

DIAGNOsIS. As for genus.

DESCRIPTION. Tests are oval in outline with a rounded
anterior and slightly pointed and projecting posterior (Pl. 23,
figs 1-3). In profile the test is low domal with a rounded
ambitus and slightly angled posterior (PI. 23, fig. 4). The oral
surface is concave towards the peristome. Test length ranges
from 55 to 95 mm. Test width is 65-81% of test length (mean
= 76%, N = 15), test height 35-47% of test length (mean =
40%).

The apical disc lies 37-44% of test length from the anterior
border. It is tetrabasal (Fig. 62), with genital plates 1, 2 and 4
reduced to minute plates largely occupied by the gonopores.
The madreporite is stellate in form.

Petals are long and weakly lanceolate in outline (Fig. 61).
The anterior petal and the two posterior petals are similar in
length, being 35-42% of the test length. The latero-anterior
petals are shorter, typically 80-90% of the length of the other

8 Ambulacrum
@ Interambulacrum

30 40 50 60
Test diameter (mm)
70 a
gs Test width 5
@ Test height i Les

60 70 80 90 100
Test length (mm)

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 201

Were

pe BRS,
eee |

PLATE 23

' Figs 1-4, 7 Pygurostoma morgani Cotteau & Gauthier. 1, specimen in the Morgan Collection, Museum d’Histoire Naturelle, Paris; oral, x
1. Senonian of Iran; no locality details. 24, 7, BMNH EE3315; 2, apical, x 0-9; 3, oral, x 1; 4, lateral, x 1; 7, detail of peristome x 6.
Jebel Rawdah, section 2, loose in scree at level of bed 12.

| Figs5,6 Petalobrissus linguiformis (Peron & Gauthier). BMNH EE4318; 5, apical; 6, lateral; both x 2. Jebel Rawdah, section 2, bed 21.

Mere

|
AANA

=:
>=

yp :
ay

Hips,
Br.
Z
Z

; 1
Ba a ai
~. Ps
ZZ, 5 2
MMi STATA

Pa
i”

WWYE™
oat
i

\\\

SS x

Fig. 61
oral. Scale bar = 1 cm.

three. The anterior petal is also typically narrower and more
parallel-sided than the other petals. The two pores are
subcircular and widely separated, united by a well developed
groove. The petals converge slightly distally but do not close
(Fig. 61). The interporal zone is about twice the width of a
single pore zone. Pores below the petals are all single.

The peristome is quinquestellate in outline with the promi-
nent bourrelets projecting into the opening (PI. 23, fig. 7).
The bourrelets are triangular and have long lateral bands of
fine tuberculation. They are blunt-ended. The phyllodes are
distinctly sunken and arcuate. They consist of many rows of
pores (Fig. 63). There is a well defined outer series and inner
series in each half ambulacrum, but the central zone consists
of a broad band of unorganised pores, four or five abreast.
The anterior ambulacrum, however, has notably fewer
median pores than other ambulacra, with only one or two
median pores abreast. There are 16-18 pores in the outer
series and 7 or 8 in the inner series. There are many occluded
plates in the phyllodes (Fig. 63). The peristome is 1-4 to 2-1
times as wide as it is long and is situated 33-43% of test length
from the anterior border (mean = 39%, N = 13).

The peristome is small and lies inframarginally, typically
forming a small pointed rostrum. There is a distinct, well-
formed invagination of the test on the adoral margin of the
periproct but not adambitally. The periproct is oval and
longer than wide (width = 45-70% of length). It opens
between interambulacral plates 6 and 8 (Fig. 61).

Tuberculation is fine and dense aborally, slightly less fine
and less dense adorally. There is a very pronounced fusiform

A.B. SMITH

Camera lucida drawings of plating in Pygurostoma morgani Cotteau & Gauthier. A, BMNH EE3303, apical; B, BMNH EE3304,

naked zone along the sternum between the peristome and
periproct. This is slightly raised above the surrounding test
and is covered in small pits. There is a similar, smaller naked
zone developed anterior of the peristome in ambulacrum III.

REMARKS. The species was described and figured by Cotteau
& Gauthier (1895). In the same work they described a
number of smaller species of Parapygus under the name
Pseudocatopygus. Some of these may possibly turn out to be
juveniles of P. morgani.

This species is very different from the only other possible
species assigned to this genus, P. pasionensis Cooke (1949),
from the ?Campanian of Guatamala. P. pasionensis has much
less well developed bourrelets and its periproct is more
equant and lies completely flush and is not invaginated. As
discussed above, it is not clear that P. pasionensis is conge-
neric with P. morgani.

Family CASSIDULIDAE Agassiz & Desor, 1847

Genus PETALOBRISSUS Lambert, 1916
Petalobrissus rawdahensis sp. nov. Pl. 24, figs 1-12;
Figs 64, 65

Types. Holotype, BMNH EE3485; paratypes, BMNH
EE3467-84, EE3486-87, EE4321-22.

OTHER MATERIAL. Over 650 specimens were collected.

{

_ LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

Fig. 62 Camera lucida drawing of apical disc plating in
Pygurostoma morgani Cotteau & Gauthier, BMNH EE3288.
Scale bar = 1 mm.

‘Fig. 63 Camera lucida drawing of phyllode plating in Pygurostoma
morgani Cotteau & Gauthier, BMNH EE3304. Scale bar = 5
mm.

‘OCCURRENCE. This species occurs abundantly at Jebel Raw-
‘dah, section 2 but is found virtually nowhere else. The only
‘other occurrence of this species is in the lowest bed (bed 1) at
Jebel Rawdah, section 3b. At section 2 the species is found at
ithe following levels: bed 4 (3); bed 8 (4); bed 11 (247); bed

12/13 (230); bed 14 (17); bed 15 (25); bed 19 (73); beds 20-21

(45); bed 22 (13); bed 26 (2). Additional material was
collected loose from the scree at this locality.

DiaGNosis. A Petalobrissus with a monobasal apical disc
and a small quinquestellate peristome, anterior in position
and surrounded by small pointed bourrelets. Phyllodes well-

203

developed, composed of outer and inner series of pores.
Buccal pores present. Periproct posterior, longitudinal. Nar-
row, smooth tubercle-free zone present both anterior and
posterior to the peristome. In outline the posterior projects
slightly and adorally there is a characteristic interradial keel
in the posterior interambulacrum.

DESCRIPTION. Tests are oval in outline, uniformly rounded
at the anterior, but slightly pointed posteriorly (Pl. 24, figs 5,
6). Test length ranges from 10-3 to 21-2 mm. Test width is
88-95% of test length (mean = 91%, SD = 1:8%, N = 23)
and the widest point is slightly posterior of midlength. In
profile the test is depressed with a relatively flat upper
surface, a uniformly rounded anterior and an obliquely
truncated posterior (PI. 24, figs 4, 7, 10, 11). The tallest point
coincides with the apical disc or is slightly posterior to it. The
ambitus is rounded and relatively low. Test height is 51-62%
of test length (mean = 56%, SD = 2-4%, N = 23; Fig. 64).
The oral surface is flat anteriorly, but slopes away towards
the posterior. There is a relatively sharp and well-defined
posterior keel on the oral surface along the midline and two
less sharply defined keels in the two postero-lateral interam-
bulacra (PI. 24, figs 5, 8).

The apical disc lies 37-46% of test length from the anterior
border (mean = 43%, SD = 2:5%, N = 23). It is monocyclic,
with four gonopores that open at around 10-13 mm test
length. The gonopores are oval and there is no sign of any
suture separating them from the central area of madrepores
(Fig. 65C). Ocular plates are small and subtriangular in
outline.

Petals are bowed and converge distally, though remaining
open (Fig. 65A). The anterior petal is the longest and the
least bowed. It has about 20 pore-pairs in a column at test
length of 10 mm, rising to 39 at 21 mm test length. The
perradial portion of the ambulacrum is about 1-5 times the
width of a single pore zone. Lateral and posterior petals are
60-86% of the anterior petal in length and are more strongly
bowed. In all petals the two columns are of equal length. The
posterior petals end well short of the periproct.

All pores below the petals are single. The phyllodes are
well developed and strongly bowed but are only slightly
sunken (PI. 24, figs 1, 5, 12). The first ambulacral plates are
elongate and bootshaped in outline, with small buccal pores
(Fig. 65D). There is an outer series of closely packed pores
and a shorter inner series of equally closely spaced pores (PI.
24, fig. 12; Fig. 65D). There are 10 or 11 pores in the outer
series of lateral and posterior phyllodes, and 9 or 10 in
ambulacrum III. There are 4 or 5 pores in the inner series in
all ambulacra. Despite the large number of individuals avail-
able, none show the detailed plating of the phyllode region
adequately. The outer series of pores lie on a series of narrow
plates that do not reach the perradius. Internally there is a
second series of plates housing the inner series of pores.
These may be oblique, but in no specimen is the plating in
this region clear.

The peristome is pentagonal, slightly broader than long in
larger individuals, and situated 33-39% test length from the
anterior border (mean = 36%, SD = 1:5%, N = 23). The
peristome width is 7-12% of test length and its length is
85-100% of its width. The peristome is sunken with well
developed vertical walls. The surrounding interambulacral
areas are developed into short, projecting, knob-like bourre-
lets, which do not, however, impinge on the peristome (PI.
24, fig. 12).

204 A.B. SMITH

PLATE 24

Figs 1-12 Petalobrissus rawdahensis sp. nov. 1-4, 12, BMNH EE4321, holotype; 1, oral; 2, apical; 3, posterior; 4, lateral; all x 3; 12, detail |
of peristomal region, x 6. Jebel Rawdah, section 2, loose in scree at level of bed 11. 5-7, BMNH EE3485, paratype; 5, oral; 6, apical;7, |
lateral; all x 3. Jebel Rawdah, section 2, bed 11. 8-11, BMNH EE4322, paratype; 8, oral; 9, apical; 10, lateral; 11, posterior; all x 3. Jebel |
Rawdah, section 2, loose in scree at level of bed 11.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

E @ P. rawdahenss: to periproct
£ @ P. cf. setifensis: to apical disc
- © P. cf. setifensis: to periproct
_
fe} @
20 i
©
cd
s oa
rs °
© mR .
Se 1@) oe [ao] oO
©
o La Pe ad =
s ed
= e
a
ac)

10 20 30
Test length (mm)

= i)
(-) co [o} N

To peristome from anterior (mm)
bd

20 30
Test length (mm)

+
fo)

@ P. rawdahensis o
@ P. cf. setifensis

Ww
(“al

Ww
[o)

Number of pore-pairs in column
N
on

4 6 8
Length of petal in ambulacrum Ill

205

© P. rawdahensis: test width
@ P. rawdahensis: test height
@ P. cf. setifensis: test width
© P. cf. setifensis: test height

10 20 30 40
Test diameter (mm)

es P. rawdahensis
@ P. cf. setifensis

Test length (mm)

40
@ P. rawdahensis
@ P. cf. setifensis

3 4 5 6 7 8
Length of petal in ambulacrum Il

Fig. 64 Biometric data for Petalobrissus rawdahensis sp. nov. and P. cf. setifensis (Cotteau).

The periproct is clearly visible from above and opens
71-87% test length from the anterior border (mean = 82%,
SD = 2:7%, N = 23). It is longitudinal, with a width that is
41-65% of its height. The opening is near vertical, with
slightly invaginated walls forming a short, parallel-sided anal
sulcus. There is a subanal rostrum (PI. 24, figs 2, 6, 9-11).

Tuberculation is fine and uniform aborally, slightly coarser
adorally. On the oral surface there is a narrow band free of
tubercles down the midline of the posterior interambulacrum.
This is finely granular and tapers towards the posterior.

‘REMARKS. This species is problematic to place on account of
its monobasal apical disc. In general shape and plating it
conforms closely to species of Petalobrissus, having very
‘similar phyllodes and bourrelets, a similar longitudinal

/Periproct set far back on the test, and only a remnant anal

sulcus. However, the type species of Petalobrissus has a
tetrabasal apical disc according to Kier (1962), whereas P.
rawdahensis has a monobasal disc. P. rawdahensis also bears
a strong resemblance to small Hardouinia species, especially
in the characteristic keeled oral surface, bowed petals and
well developed bourrelets. However, in the type species of
Hardouinia, H. mortonis, the phyllodes are more strongly
arcuate with the inner series arranged distally as an integral
part of the arc. Furthermore, the peristome opens subcen-
trally. In the type species of Procassidulus, P. lapiscancri
(Goldfuss) from the Maastrichtian of Maastricht, the phyl-
lodes are less arcuate and the inner series is parallel to the
outer series of pores. P. rawdahensis also differs from P.
lapiscancri (Goldfuss) in being larger, more rounded in
profile and oval in outline, and in having more pores in its

206

A.B. SMITH

Fig. 65

Camera lucida drawings of plating in Petalobrissus rawdahensis sp. nov. A, apical surface, BMNH EE3485; B, oral surface, BMNH

EE3487; C, apical disc, BMNH EE3476; D, One phyllode, peristome margin at base, BMNH EE3484. Scale bars = 1 mm.

phyllodes at comparable sizes. Pending revision of this group,
P. rawdahensis is placed in the genus Petalobrissus.
Petalobrissus rawdahensis is easily distinguished from Peta-
lobrissus cf. setifensis (Peron & Gauthier), which occurs in
the same section, by its more angular outline, its posterior
keel and slope on the oral surface, its smaller, less quinquelo-
bate peristome and its more bowed and densely pored petals.

Petalobrissus cf. setifensis (Cotteau, 1866) al, 25);
figs 1-10; Figs 64, 66

cf. 1866 Echinobrissus setifensis Cotteau: 267, pl. 14, figs
13-15.
cf. 1962 Petalobrissus setifensis (Cotteau); Kier: 125, pl. 16,
figs 10-13.
1989 Petalobrissus inflatus Gauthier & Thomas; Ali:

405, fig. 5 (9).

MATERIAL STUDIED. Thirty one specimens were used for the
biometric analysis: BMNH EE3505-10, EE3512-28, EE3530,
EE3532, EE3535, EE3537-41. An additional 269 specimens
were collected.

OCCURRENCE. Apart from five specimens collected from the
scree at Jebel Buhays, section 1, and a single specimen from
bed 9 at Jebel Bu Milh, all the material comes from sections

at Jebel Rawdah. The distribution of specimens is as follows:

Jebel Rawdah, section 1: bed 3 (18).

Jebel Rawdah, section 2: bed 8 (9); bed 11 (124); beds 12/13
(4); bed 14 (15); bed 19 (23); bed 21 (44); bed 26 (3).

Jebel Rawdah, section 3b: bed 2 (4).

Jebel Rawdah, section 4: bed 4 (1); beds 8/9 (3).

DESCRIPTION. Tests are oval in outline, uniformly rounded
at the anterior and slightly flattened at the posterior (PI. 25,
figs 1, 2). Test length ranges from 10 to 33 mm. Test width is
82-93% of test length (mean = 87%, SD = 2-:9%, N = 3}),
with the widest point slightly posterior of midlength. Test
height is 49-61% of test length (mean = 54%, SD = 3-1%,N
= 31), with the highest point at or a little posterior of the
apical disc. In profile the anterior is uniformly rounded, the
posterior obliquely truncated (PI. 25, figs 4, 10). The ambitus
is rounded and about one-quarter of the test height above the
base. The oral surface is flat, with a very slight depression
towards the peristome.

The apical disc lies 37-48% test length from the anterior
border (mean = 42%, SD = 2-4%, N = 31) being relatively
more anterior in larger individuals (Fig. 64). It is monocyclic
with no evidence of any sutures between the gonopores and
central madreporite region in any specimen, even where
ocular plate sutures are clear (Fig. 66D). However, the

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 207

PLATE 25

Figs 1-10 Petalobrissus cf. setifensis (Cotteau). 1, 10, BMNH EE4341; 1, oral; 10, lateral; both x 3 Jebel Buhays, section 1; loose in the

_ scree derived from the lowest 3 m of the Simsima Formation. 2-5, BMNH EE3519; 2, apical; 3, oral; 4, lateral; 5, posterior; all x 3. Jebel
Rawdah, section 2, bed 11. 6-9, BMNH EE3536; 6, apical; 7, oral; 8, lateral; 9, posterior; all x 3. Jebel Rawdah, section 2, bed 19.

Figs 11,12 Arnaudaster cylindriformis sp. nov. BMNH EE4324, depressed variety; 11, oral; 12, apical; both x 2 (see also Pl. 29, Figs 6,
9). Jebel Rawdah, section 2, in scree at level of bed 14.

208

A.B. SMITH

Fig. 66 Camera lucida drawings of plating in Petalobrissus cf. setifensis (Cotteau). A, apical surface, BMNH EE3514; B, oral surface,
BMNH EB3535; C, phyllode plating, BMNH EE3524; D, apical disc, BMNH EE3524; E, single phyllode, peristome margin to base,

BMNH EE3535. Scale bars: A, B = 2 mm, C-E = 1 mm.

madrepores do not usually extend up to the gonopores.
Gonopore diameter varies markedly between individuals of
the same test length, indicating sexual dimorphism in this
species. Gonopores appear at around 10-12 mm test length.
Petals are lanceolate, with almost straight lines of inner
pores and only slightly bowed lines of outer pores in all petals
(Pl. 25, fig. 2; Fig. 66A). All are distally open and both inner

and outer pores are subcircular in outline. The anterior petal
is the longest and has about 22 pore-pairs in a column at 15
mm test length, rising to 39 at 33 mm test length. Petals in
ambulacra II and IV are 65-93% of the length of the anterior
petal (mean = 80%, SD = 6:6%, N = 28), whereas the
posterior petals are 73-100% of the length of the anterior
petal (mean = 87%, SD = 6:6%, N = 28). The posterior

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

petals end well short of the periproct.

All pores below the petals are single. The phyllodes are
only slightly expanded and are flush with the surrounding test
(Pl. 25, figs 1, 3). Irrespective of size there are 7 or 8 pores
forming an outer series and 3 or 4 pores forming an inner
series in the phyllodes (Figs 66C, E). The first ambulacral
plates are boot-shaped and carry small buccal pores, but are
not particularly elongate. The inner series of pores are found
on small occluded plates which are separated from one
another (i.e. they do not form a continuous inner series of
plates). Each is separated from its neighbours by two of the
outer plates. Sphaeridial pits are confined to the area
between the most adoral of the inner series of pores and the
buccal pores. There are just two or three in each column.

The peristome is quinquelobate, and relatively large (Fig.
66C). Its length is 9-16% of the test length (mean = 11%, SD
= 1-6%, N = 29) and 81-120% of its width. There is a distinct
vertical-walled well to the peristome, covered in fine miliary
tubercules. Bourrelets are not developed. However, the
interambulacral margins to the peristomial well are elevated
slightly, forming a lip-like rim (PI. 25, fig. 1).

The periproct is longitudinal and clearly visible from above
(Pl. 25, figs 2, 6). It lies 76-92% of test length from the
anterior border (mean = 85%, SD = 3:5%, N = 31). It is
about twice as tall as wide (mean width = 48% of height).
There is a short parallel-sided anal sulcus that stops short of
the ambitus.

All tubercles are sunken, those adorally being noticeably
larger than the aboral ones. There is a median naked zone on

| the oral surface in both the anterior ambulacrum and the

posterior interambulacrum (PI. 25, figs 1, 3, 7). The anterior

zone is short and does not reach the ambitus, but the

posterior zone is broader, approximately parallel-sided and
reaches the posterior border. It is finely granular.

REMARKS. This species could at first glance be confused with
Petalobrissus rawdahensis, but is distinguished on several
counts. It has a larger, more quinquelobate peristome, a
flatter base without the posterior keel, it lacks bourrelets and
has less inflated phyllodes with fewer pores. It has more
parallel-sided petals with less densely packed pore-pairs. It

_ resembles certain other species of Petalobrissus in its overall
| shape and form, but is distinguished by its monobasal apical

disc. In particular it most closely resembles P. cubensis
(Weisbord), particularly in its floscelle structure. However, it

| differs from P. cubensis in having a narrower posterior naked

| zone on its oral surface, in having its apical disc less anterior
and in having a proportionally longer anterior petal.

It also comes close to certain species currently assigned to
Rhynchopygus in test shape and floscelle structure, but
differs in having a longitudinal rather than a transverse
| peristome. It is closest to P. setifensis (Peron & Gauthier),
from the Maastrichtian of Algeria, but differs in some minor
details, having a flatter base and more quadrate outline with
| less rounded ambitus, and in having fewer inner-series pores.

__ This species was identified by Ali (1989) as Petalobrissus
_inflatus (Gauthier & Thomas), but P. inflatus is a Cenoma-
| nian species that differs in several respects, notably in having
“much longer and more densely pored petals at equivalent
' sizes. These petals are strongly bowed in P. inflatus and the
posterior pair extend back to a point level with the opening of
the periproct.

Cassidulus oldhamianus Stolizcka, from the Maastrichtian
Arrialoor Group of Southern India may be comparable, and

209

appears similar in overall form. However, that species is
inadequately described and figured and no examples of it are
available for comparison. At present it is impossible to say
what genus the species belongs to.

Petalobrissus_ linguiformis (Peron & Gauthier, in
Cotteau, Peron & Gauthier, 1881) Pl. 23, figs 5, 6;
Pl. 26, figs 1-6; Figs 67, 68

Types. Syntypes are the two measured specimens in the
Peron & Gauthier collections referred to in Cotteau ef al.
(1881: 162).

MATERIAL STUDIED. Ten specimens, BMNH EE3334-35,
EE3337, EE3341-45, EE4319, EE5021, were measured for
the biometric analysis. An additional 11 specimens were also
collected.

OCCURRENCE. This species was originally described from the

Maastrichtian of Algeria, and has since been reported from

Tunisia and Egypt. Apart from one specimen collected loose

at Jebel Buhays, section 3, all specimens come from Jebel

Rawdah. It is found at the following localities and horizons:

Jebel Buhays, section 1: loose in the scree, derived from the
lowest few metres of the Simsima Formation (1).

Jebel Rawdah, section 2: bed 14 (4); bed 19 (5); bed 20 (1);
bed 21 (4); loose from upper part of section (beds 14-21)
©) he

Jebel Rawdah, section 4: loose in scree (1).

DIAGNosiIs. An elongate Petalobrissus with a narrow, almost
vertical periproct and short anal sulcus set very far towards
the posterior on the upper surface. Phyllodes are very
strongly developed, with prominent bourrelets and outer and
inner series of pores forming well-defined lines. There are up
to 20 pores in the outer series and 10 in the inner series. In
addition there are well developed rows of sphaeridial pits
down the perradius.

DESCRIPTION. Tests are ovoid in outline, uniformly rounded
at the anterior and slightly more pointed towards the poste-
rior, with a slight, but distinct, anal cleft (PI. 26, figs 1, 2).
Tests range in size from 13 to 50 mm in length. The widest
point on the test lies about two-thirds the distance from the
anterior border. Test width is 76-87% of test length (mean =
82%, SD = 46%, N = 10). In profile the test is depressed,
with a height that is 43-48% of test length (mean = 46%, SD
= 1-8%, N = 9). The upper surface is slightly domed with the
tallest point slightly posterior to the apical disc. The ambitus
is uniformly rounded and lies at about one-third test height.
The posterior is more or less truncated (PI. 23, fig. 6).

The apical disc lies 34-44% test length from the anterior
border (mean = 40%, SD = 3-0%, N = 7). It is tetrabasal,
with a large madreporite plate and three small genital plates
projecting into the interambulacra (Fig. 68D). These genital
plates are U-shaped because of the gonopore that opens
along their outer margin. Genital plates lie separated from
adjacent ocular plates.

Petals are well developed (PI. 26, fig. 5: Fig. 68B). The
anterior and posterior pair of petals are both rather straight
and remain open distally, whereas the antero-lateral pair of
petals are more bowed and converge distally (PI. 23, fig. 5).
The perradial zone is about twice the width of a pore zone.
The anterior petal is noticeably longer than other petals and
extends more than three-quarters of the distance to the
ambitus. Petals II and IV are 61-74% of the length of the

210 A.B. SMITH

PLATE 26

Figs 1-6 Petalobrissus linguiformis (Peron & Gauthier). 1, 6, BMNH EE4319; 1, oral, x 3; 6, detail of peristome, x 6. Jebel Rawdah,
section 2, beds 19/20. 2-5, BMNH EE3345; 2, oral; 3, lateral; 4, posterior; 5, apical; all x 2. Jebel Rawdah, section 2, loose, derived from
beds 14—21.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

= Test width Py
@ Test height a

(mm)

20 30

Test length (mm)

Length of petals (mm)

Length (mm)

Distance (mm)

211

®B Peristome
* Bourellet e

20 30 40

Test length (mm)

@ Anterior to peristome
@ Anterior to apical disc

=
jo)

(0)
30 40 Bye) 10 20 30 40 SO
Test length (mm) Test length (mm)
Fig. 67 Biometric data for Petalobrissus linguiformis (Peron & Gauthier).

anterior petal, wheras petals I and V are 71-89% of the
length of the anterior petal. The posterior petal extends less
than two-thirds the distance from the apical disc to the
periproct.

All pores below the petals are single. The phyllodes are
well developed and bow outwards strongly towards the
peristome. They become slightly sunken towards the peris-
tome, with the buccal pores situated in a shallow depression
separated from the peristomial well (PI. 26, fig. 6). There are
two series of pores, well separated from each other and with
pores closely packed together in each row. The outer row
comprises some 17 to 20 pores, the inner series 7 to 9 (Fig.
68C). The first ambulacral plates are squat and broad, with
buccal pores situated near their adambital edge and well
separated from the peristome. Pores in the outer series are
bowed and every third plate is smaller and does not reach the
inner series of plates. These pores may be slightly offset,
marking the incipient development of a median series of
pores. The inner series of pores are situated towards the
outer edge of a double column of occluded plates. Each plate
carries a sphaeridial pit on its perradial side, there being some
8-10 sphaeridial pits forming a well defined column on either
side of the perradial suture.

The peristome is subcircular to pentagonal and is approxi-
mately as broad as it is wide. It is 69% of test length in

| diameter and lies 37-42% of test length from the anterior
| border. There are short, pointed bourrelets which project
outwards but do not impinge on the peristome (PI. 26, fig. 6).

The periproct lies close to the posterior border, and is

clearly visible from above (Pl. 26, fig. 5). It is longitudinal,

being 27-40% as wide as it is tall (Pl. 26, fig. 4). It is slightly
V-shaped, narrowing adapically, and opens into a short anal
sulcus with parallel-sided walls. The periproct is almost
vertical in orientation.

Tubercles are minute and densely packed aborally, but
coarser and less dense adorally. There is a relatively broad
naked zone in the posterior interambulacrum that runs from
the peristome to the posterior border. This is 13-18% of the
test width in width. It is covered in a reticulate pattern of pits.

REMARKS. This species is readily differentiated from other
cassiduloids described here by its highly developed phyllodes,
its almost posterior periproct and its characteristic shape. The
only species that come close are the southern Indian P.
testudo (Forbes) and P. emys (Stoliczka), both from the
Maastrichtian. Both have a similar test shape, with the
posterior slit-like periproct, the anal notch and the projecting
bourrelets. P. testudo has much less developed phyllodes with
few pores in the inner series and only 8-10 pores in its outer
series. P. emys is too poorly known to be compared closely as
the original figures and description omit many important
details such as phyllode structure. It may eventually prove to
be synonymous with P. testudo.

ANY
i

A.B. SMITH

Fig. 68 Camera lucida drawings of plating in Petalobrissus linguiformis (Peron & Gauthier). A, oral surface, BMNH EE3344; B, apical
surface, BMNH EE3345; C, oral phyllode, peristomial margin at base, BMNH EE3334; D, apical disc, BMNH EE3345. Scale bars: A, B =

5mm; C, D = 1 mm.

Genus STIGMATOPYGUS d@ Orbigny, 1856

Stigmatopygus pulchellus? sp. nov. Pl. 27, figs 1-8; Figs
69, 70

Types. Holotype BMNH EE4314, paratypes, BMNH
EE3324-25, EE3329-30, EE3332-33, EE4312-13: all nine
specimens were used for the biometric analysis given below in
the description.

OTHER MATERIAL. In addition a further six specimens were
collected.

OCCURRENCE. The species has been found only at Jebel
Rawdah, section 2, at the following horizons: bed 14 (6); bed
19 (1); bed 21 (7); loose in scree, derived from beds 14-21

(1).

DESCRIPTION. Tests are elongate oval in outline, with a
rounded anterior and a small posterior indentation (Pl. 28,
figs 2-7). Tests range in length from 37 to 68 mm. Test width
is 77-84% of test length (mean = 80%, SD = 1:9%, N = 9)
and the widest point is slightly posterior of midlength. Test
height is 41-56% of test length (mean = 47, SD = 4.9%, N =
9). The test has a depressed profile with the highest point

coincidental with the apical disc and thus anterior of centre.
There is a posterior notch marking the position of the
periproct, and a small heel underneath the periproct in
profile (Pl. 27, fig. 5).

The apical disc lies anterior of centre, some 32-40% of test
length from the anterior border (mean = 37%, SD = 3-4%,
N = 7). It appears to be tetrabasal, although the madrepores
extend almost up to the start of the gonopores (Fig. 70D).
There is no evidence for sexual dimorphism amongst the
specimens to hand.

The petals are bowed, and are widest about one third of
their distance from the apex (PI. 27, fig. 7). The anterior petal
has columns of equal length and the inner series of pores are
almost straight. It is 22-27% of test length in length. The
interporal zone is about twice as wide as the pore zone. Petal
III is significantly longer than the rest. The petals in ambu-
lacra II and IV are bowed and converge distally but remain
open. The pore columns in the posterior petals are signifi-
cantly different in length, with the inner series some 8-10
pore-pairs shorter than the outer column in larger specimens.
These petals are also the narrowest.

All pores below the petals are single. Towards the peris-
tome phyllodes are well-developed, and expand as an open

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

213

|PLATE 27

'Figs 1-8 Stigmatopygus? pulchellus sp. nov. 14, 8, BMNH EE4314, holotype; i, lateral, x 1; 2, oral, x 1; 3, apical, x 1; 4, posterior, x 1;

8, detail of peristomial region, x 5. Jebel Rawdah, section 2, bed 19. 5-7, BMNH EE4313, paratype; 5, lateral; 6, posterior; 7, apical; all
| 2. Jebel Rawdah, section 2, bed 14.

/
|
|
|

214
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Test length (mm)
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i=
©
=

Test length (mm)

Fig. 69 Biometric data for Stigmatopygus pulchellus sp. nov.

‘V’ (Pl. 27, fig. 8). There are five or six pores in each series
along the margin with an additional one, or rarely two inner
pores at the distal end of the phyllodes (Fig. 70C). Usually
there is only one occluded plate in each phyllode, all other
plates extending from the adradial to the perradial suture.
The first ambulacral plates are long and boot-shaped, with
buccal pores that are much smaller than those composing the
remainder of the phyllodes (Fig. 70C). Sphaeridial pits occur
on each plate close to the midline, forming an alternating
series, 4 or 5 per column. The phyllodes lie sunken relative to
the surrounding test with buccal pores on the adoral walls of
this depression, rather than in the peristomial well.

The peristome is pentagonal and either equilateral or
slightly longer than wide (PI. 27, fig. 8). Bourrelets are well
developed, being straight-sided to weakly wedge-shaped in

Anterior to peristome (mm)

. SMITH

N
a

50 60 70

Test length (mm)

40 50 60 70

Test length (mm)

outline and projecting upwards strongly. They do not, how-
ever, project over the peristome.

The periproct is only just visible from above, being situated
more or less posteriorly (Pl. 27, figs 6, 7). It is triangular in
shape with an upper pinched portion and a broader, more
rounded lower portion. It is typically slightly taller than
broad. There is a subanal platform that extends as an
invaginated floor to the periproct. There is a slight aboral lip
to the periproct. It lies 21-34% above the base of the test
(mean = 27%, SD = 5-5%, N = 7).

Tuberculation is fine and uniform aborally, slightly coarser
orally. There is a very broad and well-developed naked zone
running the length of the posterior interambulacrum. A
smaller naked zone is also present anteriorly along the
midline. These zones are covered in a very fine granulation

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

| =1mm.
but appear smooth to the naked eye.

' REMARKS. This is a very distinct species, on account of its
/ asymmetric posterior petals and keyhole-shaped periproct.
Small forms resemble Petalobrissus linguiformis (Peron &
Gauthier), but are easily distinguished from that species by
| their phyllode structure and periproct shape. P. linguiformis
has a longitudinal periproct with a short parallel-sided anal
sulcus, and also has much better developed phyllodes, with a
separate series of inner occluded plates.

The new species undoubtedly comes closest to the type
species of Stigmatopygus, S. galeatus d’Orbigny from the late
Cretaceous of Angouleme, France. Both have a very similar
test shape and periproct position and shape. Unfortunately,
this species is very poorly known, the original description and
| figures being quite inadequate by today’s standards. In par-
ticular its phyllode structure is unreported. Kier (1962)
described the phyllode structure of another species S. lam-
berti Bessairie, from the Campanian of Madagascar, but this
\differs in being much wider and taller, with a much larger
periproct. It also has better developed phyllodes, which are
clearly bowed and comprise an outer series of some 12 pores

i)
—
Nn

| Fig. 70 Camera lucida drawings of plating in Stigmatopygus pulchellus sp. nov. A, oral surface, BMNH EE4312; B, apical surface, BMNH
EE4314; C, phyllode plating, peristomial margin at base, BMNH EE4312; D, apical disc, BMNH EE3325. Scale bars: A, B = 5mm; C, D

and an inner series of 4 or 5 pores. I am therefore not certain
that S. /Jamberti is truly congeneric with the type species S.
galeatus. Only re-study of the type (apparently lost), or
topotype material will solve the problem. If the phyllode
structure of S. galeatus is similar to that of S. pulchellus, then
S. lamberti should be transferred to a new genus. On the
other hand, if S. galeatus proves to have a phyllode structure
similar to that of S. lamberti, then S. pulchellus should be
made the type of a new genus. Consequently, S. pulchellus
can only tentatively be placed in the genus Stigmatopygus.

S. galeatus can be distinguished from S. pulchellus by the
fact that its petals are illustrated as being of equal length.

216

ws Test wdth
@ Test height

(mm)

30 40
Test length (mm)

gs Amb. lll
@ Amb. Il r
O Amb. | L

—)
jo)

fee}

Petal length (mm)

30 40

Test length (mm)

Peristome width (mm)

1 2 3
Peristome length (mm)

Fig. 71 Biometric data for Nucleopygus magnus sp. nov.

Genus NUCLEOPYGUS Agassiz, 1840
Nucleopygus magnus sp. nov. PI. 28, figs 1-7; Figs 71,
2

Types. Holotype BMNH EE4339, paratypes, BMNH
EE3340, EE3356, EE3358, EE3363, EE3365, EE3367-68,
EE4327, EE4335-38

OTHER MATERIAL. An additional 29 specimens were col-
lected. Biometric data was taken from the type series only.

OCCURRENCE. This species was found at the following locali-

ties and horizons:

Jebel Buhays, section 1: loose in scree, derived from the
lowest few metres of the Simsima Formation (19).

Anterior to peristome (mm)

A.B. SMITH

20 30 40 50
Test length (mm)
40
ss a Amb. lll »
oO
- @ Amb. Il
& O Amb. | a
£
2
ss
a
©
ro
a
i)
h
o
2
E
=)
z
20 30 40 50
Test length (mm)
40

® to apical disc
* tostart of anal sulcus

Distance from anterior (mm)

30
Test length (mm)

40 50

Jebel Buhays, section 3: loose, derived from the lowest bed of
the Simsima Formation (1).

Jebel Thanais: lowest 2 m of the Simsima Formation (4).

Jebel Rawdah, section 2: bed 14 (8); loose in scree (3).

Jebel Rawdah, section 4: bed 12 (1).

Jebel Faiyah, section 1b: bed 2 (1).

DIAGNOsIs. A very large, elongate Nucleopygus with a deep
median depression on the oral surface. The periproct opens
relatively close to the posterior border and there is only a
short anal sulcus. The posterior petals end at a level slightly
anterior to the start of the anal sulcus.

DESCRIPTION. Tests are subquadrate in outline, with a
rounded anterior and a somewhat truncated posterior with a
shallow anal notch (PI. 28, figs 1, 4, 5). Test length ranges

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 217

& &

a a

é,
Serve rere re

*
%
*

PLATE 28

Figs 1-7 Nucleopygus magnus sp. nov. 1-3, BMNH EE4327, paratype; 1, apical, x 2; 2, lateral, x 2; 3, detail of apical disc, x 4. Jebel
Thanais, lowest 2 m of the Simsima Formation. 4-6, BMNH EE4339, holotype; 4, apical; 5, oral; 6, lateral; all x 1. Jebel Buhays, section

1; loose in the scree’derived from the lowest 3 m of the Simsima Formation. 7, BMNH EE3367; detail of peristomial region, x 4. Jebel

| Buhays, section 2; loose in the scree derived from the lowest 3 m of the Simsima Formation.
Figs 8-10 Nucleopygus iranicus (Cotteau & Gauthier). Specimen in the Morgan Collection, Museum d'Histoire Naturelle, Paris; 8, lateral;

9, apical; 10, oral; all x 2. Senonian, Poucht-e-Kouh, Iran.

218

A.B. SMITH

Fig. 72 Camera lucida drawings of plating in Nucleopygus magnus sp. nov. A, apical surface, BMNH EE4339; B, oral surface, BMNH
EE4337; C, apical disc, BMNH EE4339; D, phyllode plating, BMNH EE4337. Scale bars: A, B = 5 mm; C, D = 1 mm.

from 18 mm to about 60 mm (estimated from a broken
specimen). Test width is 76-82% of test length (mean = 79%,
SD = 2%, N = 11), and the widest point on the test is about
two thirds of the distance back from the anterior. The test has
a low profile, with test height 42-51% of test length (mean =
39%, SD = 3%, N = 9). It is rounded towards the anterior
but obliquely truncated towards the posterior (PI. 27, figs 2,
6). The ambitus is rounded, but relatively low. The oral
surface has a marked median depression and the peristome is
sunken (PI. 28, fig. 5).

The apical disc lies 36-43% of the test length from the
anterior border (mean = 39%, SD = 1%, N = 9). It is
tetrabasal with large gonopores projecting into the interam-
bulacra (Fig. 72C). Ocular plates are small and U-shaped.
The small genital plates are separated from one another by
ocular plates.

Petals are open and subparallel to very slightly bowed
(lateral and posterior pairs) (PI. 28, figs 1, 4; Fig. 722A). Pores
are both approximately circular and are joined by a well-
marked furrow (PI. 28, fig. 3). The interporal zone is approxi-
mately 1-5 times as wide as a pore-zone. The anterior petal is
the longest, but is only slightly longer than the posterior
petals. It has 30 pore-pairs in a column at 30 mm test length,

rising to 40 at 40 mm test length. The posterior petals end
slightly in front of the anal sulcus.

All pores are single below the petals. Phyllodes are hardly
expanded adorally (PI. 28, fig. 7; Fig. 72D). There are buccal
pores at the rim of the peristome, situated on relatively short
and squat first ambulacral plates. The phyllodes have both an
outer and an inner series of pores, although the two series are
not well separated (Fig. 72D). There are six pores per column

in the outer series in ambulacrum III with 7 or 8 in the other |

ambulacra, and | or 2 pores in the inner series of ambulacrum
III with 2 or 3 in other ambulacra. Every third ambulacral
plate is occluded, but the occluded plates do not form a
continuous inner series. The phyllodes are not depressed
relative to the surrounding test. There are no bourrelets, but
the walls to the peristome are covered with fine, dense
tubercules. The posterior interambulacrum differs from other
interambulacra in that it does not turn sharply in towards the
peristome, but instead forms a sloping shelf into the opening.

The peristome is pentagonal in outline, approximately 1-6
times as wide as long. It lies 38-42% of the test length from
the anterior border (mean = 40%, SD = 1%, N = 10).

The periproct is tall and narrow and has a width approxi-
mately 40% of its height. It opens into a narrow, parallel-

:
|

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

sided anal sulcus that slopes posteriorly. The periproct opens
73-85% of test length from the anterior border (mean =
78%, SD = 4%, N = 11).

The upper surface is covered in fine, slightly sunken

~ tubercles. Adorally tubercles are coarser but are still sunken.
_ There is a narrow, zig-zagged granular zone in the posterior
-interambulacrum that follows the interradial suture and

tapers out before reaching the pesterior border (PI. 27, fig.
7). It is never more than about 78% of the test width at its

' widest.

REMARKS. Few other species of Nucleopygus come any-

_ where near the size of N. magnus. Both N. iranicus (Cotteau

}

& Gauthier) (Pl. 28, figs 8-10), from the late Cretaceous of
southern Iran, and N. pullatus Stolizcka, from the Maastrich-
tian of southern India are much smaller species, reaching
little more than 14 mm in test length. Furthermore, they are
squatter. N. geayi (Cottreau), from the Maastrichtian of
Madagascar, is even smaller, never being reported larger
' than 8 mm test length. Similarly, the European species N.
. coravium and N. scrobiculatus are equally small forms.
Only in the late Cretaceous of North Africa do any
_ Nucleopygus approach the size of N. magnus. N. inaequalis
(Peron & Gauthier) resembles N. magnus in shape but its
posterior petals extend posteriorly beyond the start of the
_ anal sulcus. N. meslei (Peron & Gauthier), from the Campa-
| nian, is probably the closest. N. meslei reaches 30 mm in test
‘length, but differs from N. magnus in having a more anterior
_ apical disc, a longer anal sulcus and more petaloid ambulacra.
/ It is also less markedly depressed along the midline on the
_ oral surface.

Family ECHINOLAMPADIDAE Gray, 1851
Genus ARNAUDASTER Lambert, 1918

REMARKS. Until now there has only been one species
assigned to this genus, A. gauthieri Lambert, from the
_Cenomanian of France (Aquitaine). Kier (1962: 105) noted
that Arnaudaster might best be considered a synonym of
Parapygus, as it differs only in having a more cylindrical
shape and more unequal poriferous zones in the same petal.
| The discovery of a Maastrichtian species very close in form to
_A. gauthieri supports the maintenance of Arnaudaster as a
separate genus. Pseudocatopygus longior Cotteau &
Gauthier, from the late Cretaceous of Iran, also has unequal
columns in its petals and may belong to Arnaudaster. How-
ever, Kier (1962) placed Pseudocatopygus as a synonym of
_ Parapygus.

_Arnaudaster cylindriformis sp. nov. PI. 25, figs 11, 12;
Pl. 29, figs 1-9; Figs 73-75

Types. The holotype is BMNH EE4334, paratypes are
|BMNH EE4324, EE4331-33, EE3378-81.

OTHER MATERIAL. Three poorly preserved specimens,
BMNH EE3374-76. BMNH EE3373 may also belong here,
but only a fragment of the upper surface is preserved and this
apparently shows equally developed columns of pore-pairs in
-ambulacrum petal V.

OCCURRENCE. This species was found at the following locali-
ties along the western margin of the Oman Mountains:
Jebel Buhays, section 1: top of bed 1 (1); loose in the scree,

219

ws Test width (mm)
@ Test height (mm)

Test length (mm)

B To peristome oO
@ To apical disc

=
a

Distance from anterior (mm)
N

10
8
6
20 30 40 50 60
Test length (mm)
20
a Ambulacrum iil
=e @ Ambulacrum Il JN
£ A Ambulacrum!
= 15 ve =
= A 5 °
a A er ¢
- A a °
010 ] r :
€ oe?
= °
®
—_

20 30 40 50 60
Test length (mm)

Fig. 73 Biometric data for Arnaudaster cylindriformis sp. nov.

derived from the lowest few metres of the Simsima Forma-
tion (6).

Jebel Rawdah, section 2: bed 14 (1); loose at the level of bed
19 (2); loose in scree, derived from beds 14~21 (2); bed 21
(1); bed 26 (1).

Jebel Rawdah, section 3: bed 9 (1); loose near top of section

(1).

DIAGNOsIS. An Arnaudaster with a wide, pentagonal peris-
tome with weak bourrelets and approximately four pores in
the inner series of phyllodes in each half ambulacrum.

DESCRIPTION. Tests range from 21 to approximately 50 mm
in length, although many specimens are lacking the very

720 A.B. SMITH

PLATE 29 |
Figs 1-9 Arnaudaster cylindriformis sp. nov.14, BMNH EE4334, holotype; 1, oral, x 2; 2, apical, x 2; 3, detail of peristomial region, x 4, |
4, lateral, x 2. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation. 5, 7, 8, BMNH EE4323;,

5, apical; 7, lateral; 8, oral; all x 2. Jebel Rawdah, section 2, bed 21. 6, 9, BMNH EE4324 (depressed variety); 6, posterior; 9, lateral; both |
x 2 (see also Pl. 25, Figs 11, 12). Jebel Rawdah, section 2, bed 14.

| LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 221

at ACN,
PAARL

ua
=

catty
, My
“Ny ly
Uy My A ww
My ny HAWN

Fig. 74 Camera lucida drawing of plating in Arnaudaster cylindriformis sp. nov, A, oral surface, BMNH EE4324; B, apical surface, BMNH

EE4333. Scale bar = 5 mm.

Fig. 75 Camera lucida drawings of plating in Arnaudaster cylindriformis sp. nov. A, apical disc, BMNH EE3379; B, phyllode plating,
BMNH EE4324. Scale bars = 1 mm.

222

posterior end of the test. Tests are ovoid in outline with a
rounded anterior and a very weakly pointed posterior (PI. 25,
figs 11, 12; Pl. 29, figs 1, 8). Test width is 71-78% of test
length (mean = 74%, SD = 2:5%, N = 10), with the widest
point on the test approximately two thirds of the distance
from the anterior and well behind the level of the apical disc.
Test height is 41-60% of the length (mean = 51%, SD =
6:1%, N = 10). In profile the upper surface is broad and
almost flat, uniformly rounded at the anterior, but undercut
at the posterior (PI. 29, figs 4, 7, 9). The oral surface is flat or
very slightly sunken towards the peristome, while the ambitus
is smoothly rounded.

The apical disc lies well anterior of centre, some 31-41% of
test length from the anterior border (mean = 37%, SD =
2:9%, N = 8). Plating is tetrabasal, although genital plate 2 is
enormously enlarged in comparison to the other genital
plates and occupies the centre of the disc (Fig. 74A). The
entire madreporic plate is covered in dense, fine perforations.
Other genital plates are very small and are dominated by the
gonopores. There may be sexual dimorphism in the size of
gonopore openings. Ocular plates are small, approximately
the same size as the genital plates.

Ambulacra are petaloid with the interporal zone slightly
inflated. The anterior and two posterior petals are similar in
size, but the lateral petals are shorter. Petals are broad,
lanceolate with more or less parallel columns of pore-pairs at
the distal tip (Pl. 29, fig. 2; Fig. 74B). They extend about 80%
of the distance to the ambitus. Pore-pairs are conjugate. The
anterior petal has columns of equal length, with 41 pore-pairs
in a column at 32 mm test length, rising to 57 at 50 mm test
length. The anterior column of pore-pairs in the lateral petals
is shorter than the posterior column by about five pore-pairs
(Fig. 74B). Distally they are slightly narrower, but remain
broadly open. The posterior petals also have unequally
developed columns of pore-pairs, with the posterior column
shorter than the anterior column by seven or so pore-pairs.
They extend approximately 70% of the distance to the
ambitus and remain broadly open distally.

Below the petals pores are single. Phyllodes are not
strongly developed and phyllode plating is best seen in
BMNH EE3379. Pores of the outer series become enlarged
close to the peristome, but the rows of pores do not bow out
(Fig. 75B). There are only eight or nine pores forming the
outer series in one half ambulacrum. There are three or four
pores forming an inner series in each half ambulacrum. Each
inner series pore is found on an occluded plate (Fig. 75B).
There are small but obvious buccal pores which are not
separated from the other pores of the phyllode but which
open right at the peristomial rim.

The first interambulacral plates are short and wide and are
slightly swollen to form a weak floscelle (Pl. 29, fig. 3; Fig.
75B). The peristome is pentagonal, approximately twice as
wide as long, and with well-developed vertical walls.

The periproct is small and transversely oval, with a width
approximately 50-70% of its height. It lies at approximately
mid-height on the posterior face, but is angled slightly
downwards so that it is just visible from the oral surface and
not from the adapical surface (PI. 29, figs 5, 8). Because the
posterior of the test is slightly drawn out, the periproctal
region is often damaged. The base of the periproct lies
between about 30 and 40% of the test height above the base.
Tubercles are small, sunken and densely crowded over the
whole surface.

A.B. SMITH

REMARKS. This species comes very close to the type species |

A. gauthieri in almost every respect. Both have a very similar,

cylindrical test, anterior apical disc and strongly unequal | |

same columns that are asymmetric). The only difference of |

columns of pores in both lateral and posterior petals (the |

significance appears to be in the development of phyllodes. In |

A. gauthieri the inner series of pores is rather poorly devel-

oped and not well separated from the outer series. In A. | |

cylindriformis the inner series pores are somewhat more
numerous and form a distinct series. However, this is a minor
difference, and there is very little else to separate the two
forms.

The species also resembles Parapygus inflatus Cotteau & |
Gauthier in its overall shape and appearance. However,
Cotteau & Gauthier catagorically state (1895: 55) that F |

columns of pore-pairs in the petals of the species are equally
developed.

Order HOLASTEROIDA Durham & Melville, 1957
Family HOLASTERIDAE Pictet, 1857
Genus HEMIPNEUSTES Agassiz, 1836

DIAGNOsIS. Heart-shaped holasteroids showing pronounced

petal asymmetry, with the posterior column of pores well)
developed and anterior column rudimentary. Periproct sub- |

ambital and clearly visible in oral view because of the distinct |
posterior notch. The madrepores extend over all genital)

plates wedge-shaped, each just reaching the opposite adradial
suture and thus biserially arranged. |

OCCURRENCE. Late Campanian and Maastrichtian, Europe,
North Africa and the Middle East.

REMARKS. Hemipneustes is very close in appearance to Car-.
diaster. Both have a cordiform test with a deep, well-defined

anterior sulcus which has enlarged primary tubercles along its|_
border. Both genera also have a similar style of plastron) ,
plating, composed of a short, broad labral plate followed by!

up to five wedge-shaped plates which more or less occupy the

full width of the plastron (eg. Ernst 1972, text-fig. 20).\

Finally, both have rather similar petals, with the anterior

|
;

!
;

column greatly reduced in comparison to the posterior col-_

umn. The primary difference between Cardiaster and Hemip-|

neustes lies in the fact that Hemipneustes has little more than)

rudimentary pore-pairs in the anterior columns of its petals,
whereas Cardiaster has small but distinctly conjugate pore-
pairs in these anterior columns. Furthermore, madrepores
are more extensively developed in Hemipneustes, extending
beyond genital plate 2 to cover genital plate 3 and ocular
plates II, III and IV.

Many species names have been erected in the past, mostly
based on relatively few specimens and often on rather poorly
preserved material. From Noetling (1897) onwards workers
have relied on two simple biometric indicators to distinguish
species of this genus; width/length and height/length ratios)
(see for example Devries 1967; Aziz & Badve 1990). How-|
ever, not only do these ratios vary ontogenetically and thus
vary according to absolute test size, but they have also rarely
been applied rigorously using large populations.

There are three species found amongst the material col-
lected from the Oman Mountains. These appear to have
some stratigraphic value. The lowest forms found at Jebel
Rawdah belong to H. persicus var. sardanyolae Vidal, and

~ LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

are found in the basal sandy facies. These have the broad
shallow anterior sulcus characteristic of the H. delettrei com-
plex, a group restricted to the late Campanian of Tunisia and
Algeria (Zhagbib-Turki 1987). Above this, or possibly par-
tially overlapping, H. arabicus is found. This species is similar
in profile and in the positioning of its apical disc, but has a
_ much narrower, deeper and more sharply defined anterior

sulcus. The first elevated Hemipneustes forms, belonging to
_ H. compressus, appear in bed 14, having more uniformly
rounded profiles. However, by the time bed 21 is reached, H.
- compressus has a much more quadrate profile with strongly
_ pinched ridges to the frontal groove. Finally, by bed 26 some
_ H. compressus appear that are very elevated and which have
developed a strong slope towards the posterior similar to the
late Maastrichtian H. striatoradiatus.

In trying to identify the Oman/United Arab Emirates species,
a number of other species have been examined and the
| following are accepted as valid:

Hemipneustes delettrei Peron & Gauthier (PI. 30, figs 9, 10),
late Campanian of Algeria. Rounded in outline and vari-
able in profile, from highly inflated (H. africanus form) to
depressed (H. delettrei form). w/l = 0-89-0-95 (mean =
0-91): h/l = 0:58-0:79 (mean =0-70) [size range 73-98
mm]. Distinguished from all other species by the large,
broad anterior ambulacral sulcus; about twice as broad as
in any other species.

Hemipneustes striatoradiatus (Leske) (Pl. 30, figs 7, 8; Fig.

| 78C), Upper Maastrichtian of the Netherlands. Rounded

in outline with a very narrow and shallow anteal sulcus.
Variable in profile from flat-topped to distinctly raised
anterior to the apical disc. Sides rather vertical giving
quadrate cross-section. Apical disc anterior of centre. w/l =
0:87-0:97, mean = 0-91: h/l = 0-61-0-80, mean 0-69 [Size
range 50-100 mm]. Distinguished from other species by (i)
its narrow, shallow anteal sulcus, (ii) rounded outline and
(iii) quadrate lateral cross-section.

_ Hemipneustes pyrenaicus Hebert (Pl. 30, figs 5, 6) — closest to

H. compressus in having a short narrow frontal groove with

a strong vertical component, anterior apical disc, and

quadrate profile. It differs in having a flat apical surface

| and in lacking adapical pinched ridges to the anterior
sulcus.

| H. persicus Cotteau & Gauthier (PI. 31, figs 1-7; Pl. 32, figs

1-4; described below).

| H. compressus Noetling (PI. 31, figs 8-11; described below).

'H. arabicus Ali (PI. 30, figs 14; described below).

In addition the following species are treated as synonyms

or rejected names, or are based on inadequate material:

_H. minor Peron & Gauthier, Upper Senonian of Iran (a small
H. persicus Peron & Gauthier).

H. oculatus Cotteau, Maastrichtian of Ciply, Belgium. This
appears to be a large variety of H. striatoradiatus, some-
what crushed but with a deeper frontal groove as illustrated
by Cotteau. The specimens referred to under this name by
van der Ham (1989) are simply rather tall and posteriorly

inclined H. striatoradiatus.

‘H. arnaudi Cotteau 1892, Upper Senonian of Dordogne. This

is very like high forms of H. compressus in having an

elevated keel to the ambulacrum. It has too wide an

anterior sulcus to be a H. striatoradiatus. Treat as a

probable synonym.

EEE

223

H. indicus Aziz & Badve 1990; Maastrichtian of S India, is
identical in profile to H. arnaudi but larger. Its deep frontal
groove and anterior apical disc makes it part of the
compressus group.

H. sardanyolae Vidal 1921, Campanian of Spain, is treated
here as a shallow grooved variety of H. persicus.

H. nicklesi Vidal 1921, Campanian of Spain, is a Hemip-
neustes sp. based on crushed and badly preserved speci-
mens that are basically indeterminate. It is probably a
synonym of H. persicus var. sardanyolae Vidal.

Spatangoides martelli Checchia-Rispoli, Maastrichtian of
Libya. Here synonymized with H. compressus Noetling.

Spatangoides tripolitanus Checchia-Rispoli, Maastrichtian of
Libya, is clearly a species of Opisopneustes. Its primary
adapical interambulacral tubercles are well developed and
it represents a valid species, differing from O. cossoni in
having a vertically positioned periproct.

Spatangoides aichinoi Checchia-Rispoli, Maastrichtian of
Libya. Here synonymized with H. persicus.

Hemipneustes batheri Lambert [=H. leymeriei Noetling]. This
is probably a variety of H. persicus, having the posterior
apical disc and oval outline with shallow and open anterior
sulcus, but it is unusually tall.

H. noetlingi Lambert was erected as a replacement name for
H. pyrenaicus of Noetling, from the Maastrichtian of
Baluchistan. However, like Devries (1967), I can find no
significant difference between Noetling’s specimens and
the type material of H. pyrenaicus and treat H. noetlingi as
a junior synonym.

Hemipneustes leymeriei Hebert has the very narrow and
shallow groove and circular outline of a H. striatoradiatus.
Topotype material from Gansec, southern France, con-
firms this.

Spatangoides tripolitanus Checchia-Rispoli is a species of
Opisopneustes.

Hemipneustes compressus Noetling, 1897 PINS iesties
8-11; Figs 76, 77, 78A

1897 Hemipneustes compressus Noetling: 34, pl. 7, figs 3, 4,

pl. 8, figs 1, 2.

1931 Spatangoides Martellii Checchia-Rispoli: 7, pl. 1, figs
3),

1967 Hemipneustes compressus Noetling; Devries: 188, pl.
6, figs 31-33.

TyPES. The syntypes are the four specimens illustrated by
Noetling and presumably in the Geological Survey of India
collections.

MATERIAL STUDIED. Thirty specimens were collected. Bio-
metric data was taken from the following 16 specimens:
BMNH_ EE3744-45, EE3748, EE4070-71, EE4073-74,
EE4076, EE4078-82, EES022-24.

OCCURRENCE. Maastrichtian of Libya, Oman and Bal-
uchistan. In the western Oman Mountains it occurs at the
following levels:

Jebel Rawdah, section 2: bed 14 (2); bed 15 (2); bed 19 (1);
beds 20/21 (17); bed 22 (1); bed 26 (2 specimens plus
fragments); bed 27 (2).

Jebel Rawdah, section 3: beds 9/10 (1).

Jebel Rawdah, section 4: loose, towards top of section (1).

DIAGNOSIS. Test rather more elongate than in other species,
typically flat-topped in side-view and arched in cross-section.

224

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A.B. SMITH
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Sulcus depth at ambitus (mm)
N

10
Sulcus width at ambitus (mm)

12 14

Fig. 76 Biometric data for species of Hemipneustes.

w/l = 80-87% of length, mean = 83%; height 54-67% of
length, mean = 60% (size range 63-83 mm). Frontal groove
narrow but moderately deep at the anterior; typically slightly
keeled. Apical disc lies well towards the anterior (distance
between anterior and apical disc = 34-44% of length: mean
= 39%) and is pinched up. Distinguished from other species
by (i) its anterior apical disc, (ii) the deep anterior groove
which is relatively short and geniculate with a strong vertical
component, (iii) being more subquadrate outline, typically
inclined from the apical disc posteriorly.

DESCRIPTION. Tests range in length from 63 to 83 mm in
length and from 53 to 69 mm in width (width = 80-87% of
length; mean = 83%, SD = 1-8%, N = 13). In outline the
test is cordate with a sharp, deep anterior sulcus, and a

truncated and even slightly indented posterior (PI. 31, figs 8,

9). The widest point on the test lies approximately mid- |
length. Test height is variable, ranging from 38 to 54 mm |
(height = 54-66% of test length: mean = 60%, SD = 3-7%, |
N = 13). In side view the upper surface varies from almost

flat to strongly vaulted, with the tallest point on the test lying |
immediately anterior of the apical disc (Pl. 31, fig. 11). The

test slopes towards the posterior which is truncated. Anterior

to the apical disc the test curves uniformly to become almost | ,
vertical. There is a sharp curve from the anterior to the more
or less flat base (PI. 31, fig. 11). In anterior profile the test
appears vaulted with rounded sides sloping up to the crest.
The anterior sulcus becomes slightly pinched towards the | ,
base and its edges are sharply delimited (PI. 31, figs 10, 11). | )

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 225

PLATE 30

Figs 1-4 Hemipneustes arabicus Ali. BMNH EE5027; 1, oral; 2, apical; 3, anterior; 4, lateral; all x 1. Jebel Rawdah, section 1, bed 4.
Figs 5,6 Hemipneustes pyrenaicus Hebert. Specimen in the Museum d’Histoire Naturelle, Paris; Maastrichtian of Montbéraud, Haute
| Garonne, France. 5, apical; 6, lateral; both x 1.
| Figs 7,8 Hemipneustes striatoradiatus (Leske). BMNH 75822; 7, apical; 8, lateral; both x 1. Maastrichtian of Maastricht, The Netherlands.

Figs 9,10 Hemipneustes delettrei africanus Peron & Gauthier. BMNH E3654; 9, apical; 10, lateral; both x 0-75. Upper Campanian of Ain
| Joutu, Algeria.

226

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A.B. SMITH

Fig. 77 Camera lucida drawings of plating in Hemipneustes compressus (Noetling), BMNH EE3745. A, apical surface; B, oral surface. Scale

bar = 5 mm.

Fig. 78 Camera lucida drawings of apical disc plating in species of
Hemipneustes. A, H. compressus (Noetling), BMNH EE3745; B,
H. arabicus, BMNH EES027; C, H. striatoradiatus (Leske),
BMNH 38630; D, H. persicus Cotteau & Gauthier, BMNH
EE3747. Scale bars = 1 mm.

The apical disc lies at or slightly in front of mid-length
(distance to the anterior border = 34-44% of test length:
mean = 39%, SD = 3-1%, N = 13). It is elongate with 4
gonopores occupying most of the area of the genital plates
(Fig. 78A). The anterior pair of genital plates are separated
from the posterior pair by the lateral ocular plates.
Madrepores extend across genital plates 2 and 3, as well as
ocular plates II, III and IV, even extending into the wall of
the gonopores in some instances. The two posterior ocular
plates are slightly larger and abut each other.

The anterior ambulacrum is non-petalloid. It is almost

flush at the apex, becomes slightly depressed towards the

anterior and then forms a deep vertical sulcus close to the
anterior (PI. 31, figs 8, 10). This groove continues up to the
peristome and is widest near the top of the anterior sulcus
becoming narrower towards the base of the sulcus. Pore-pairs
are oblique isopores, closely spaced in the upper half, but
becoming smaller and more widely spaced towards the bot-
tom of the sulcus. The margins of the sulcus are relatively
sharp, especially towards the top of the anterior sulcus, where
they may form small crests on either side. The width of the
sulcus is 13-20% of the test width (mean = 16-3%, SD =
1-86%, N = 12) and at its deepest it is about 3-5 to 4-5 mm.

The anterior petals are strongly asymmetric, with only the
posterior column well-developed (Pl. 31, fig. 8; Fig. 77).
These curve forward and, towards the tip, turn slightly down.
The inner pore is circular, the outer one elongate, and the
distance between the pairs is as wide as the outer slit-like
pore. Pore-pairs are conjugate and are separated from one
another by a single row of small tubercles. The posterior
column tapers both adapically and adambitally. There are 45
pore-pairs in a column in a 65 mm individual, rising to 52 in
an 81 mm individual (Fig. 76). The anterior column of pores
remains rudimentary throughout. All are pore-pairs, but the
pores always remain small and close together, never becom-
ing conjugate. However, they do gradually increase in size
towards the ambitus.

The posterior petals are similar to the anterior petals in
their pore-pair development and column asymmetry, with
only the posterior column bearing large conjugate pore-pairs
(Fig. 77). This column is flexed outwards and backwards and
extends only about two thirds of the distance towards the
ambitus. There are 32 pore-pairs in a column in a 65 mm
individual, rising to 42 in an 81 mm individual. Close to the
apex the two ambulacra are almost parallel, but further away
they diverge at about 90°. Pores beneath the petals remain
double, but are microscopic. Close to the mouth there are

|
|
|

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

two to three larger peribuccal pores in ambulacrum III, four
to five in ambulacra II and IV, and three to four in ambulacra
Tand V.

The interambulacra are biserial to the apex, although they
become extremely narrow, especially posteriorly. On the oral
surface there is a short, broad labral plate followed by a series
of 5 alternating triangular-shaped plates which either just or
almost reach the opposite suture (Fig. 77). Further towards
the posterior the columns become more typically biserial.

The peristome is oval, slightly more than twice as wide as
long, and lies at the base of the anterior sulcus, some 15-18%
of test length from the anterior (mean = 17%, SD = 1:2%,N
= 11). The peristome faces forward and is visible in anterior
view. The labral plate hardly indents the posterior of the
peristome.

The periproct is oval, typically about 70% as wide as tall,
and is strongly overhung so that it is visible from the oral
surface but not from the apical surface (Pl. 31, figs 8, 9). The
test beneath the periproct is indented and there are two
projections on either side. The periproct opens between
interambulacral plates 5 and 10, the lower plates being
strongly curved. The periproct lies low on the posterior
surface and the distance to the base of the periproct from the
lower surface is some 18-27% of the test height.

Tuberculation is fine and uniform throughout, except along
the iner (interambulacral) margin of the frontal sulcus
where a double or triple row of noticeably larger tubercles is
developed. There is no trace of a lateral fasciole to be seen.
Oral tubercles are slightly larger than adapical ones, but there
is no real difference in size between the tubercles of the
plastron and those of the latero-ventral regions. The oral
ambulacra appear to be tubercle-free.

REMARKS. This species is relatively common at certain levels
in Jebel Rawdah. It co-occurs with small varieties of H.
persicus but can be distinguished from that species by its
Narrower, more vertical and more sharply defined anterior
sulcus, its more anterior apical disc, its lack of adapical
primary tubercles, and its more quadrate profile. It differs
from H. arabicus, whose size range is more-or-less coinci-

(as (a9
Li /?

l eee

DOT

dent, in having the apical disc positioned anterior of centre,
and in having a more developed vertical component to the
anterior sulcus.

The stratigraphically lowest species tend to have a more
rounded profile, while higher samples become progressively
more peaked, with a stronger posterior inclination as the
region immediately in front of the apical disc becomes taller.
Thus in the beds immediately above bed 13, Jebel Rawdah,
section 2, H. compressus is rather flat and the vertical
component of the anterior sulcus is small. By bed 21 the tests
are much more quadrate in outline, with a small but obvious
peak in front of the apical disc, and by bed 26 some highly
elevated tests are found.

Hemipneustes persicus Cotteau & Gauthier, 1895 PI.
31, figs 1-7; Pl. 32, figs 14; Figs 76, 78D, 79

1895 Hemipneustes persicus Cotteau & Gauthier: 15, pl. 2,

figs 1-6.

1895 Hemipneustes minor Cotteau & Gauthier: 17, pl. 2,
figs 7-9.

1921 Hemipneustes sardanyolae Vidal: 11, pl. 2, fig. 2; pl. 3,
fig. 2.

TYPES. The syntypes are the specimens from Aftab and
Derre-i-Chahr, Iran, described by Cotteau & Gauthier (1895:
15). Although the authors stated that they had many
examples, their description appears to have been based on
only one specimen 54 mm in length, which was illustrated.
This is here designated lectotype.

MATERIAL STUDIED. There are five large individuals of this
species, BMNH EE4083, EE4084, EE4090, EE5025-—26, all
rather poorly preserved. Small individuals that appear indis-
tinguishable are relatively common and well-preserved at one
horizon at Jebel Rawdah, section 2. Biometric data is based
on the following eleven specimens: BMNH_ EE3742,
EE3746—-47, EE4091, EE4097, EE4099, EE5028-32 in addi-
tion to the five larger specimens cited above.

OCCURRENCE. In the Oman Mountains region this species

Lens
1

Fig. 79 Camera lucida drawing of plating in Hemipneustes persicus Cotteau & Gauthier. A, oral surface, BMNH EE3747; B, apical surface,

BMNH EE3746. Scale bar = 5 mm.

228

occurs at Jebel Rawdah at the following localities and hori-

zons:

Section 1: bed 4 (2).

Section 2: bed 19 (1); beds 20/21 (23); loose in scree, derived
from beds 14-21 (4).

Section 3: bed 5 (1); bed 7 (1).

Section 4: loose a little below the top of the measured section

(1).

The type series comes from the ‘Senonienne’ (probably
Maastrichtian) of Aftab and Derre-i-Chahr, southern Iran.

DIAGNOSIS. Test ovate, depressed in side-view and in cross-
section arched. Width 80-88% of length, mean = 85%:
height 42-55% of length, mean = 49% [size range 44-89 mm
test length]. Frontal groove broad and open, expanding
anteriorly, comparatively shallow at the anterior; never
keeled. Apical disc lies approximately mid-length (distance
between anterior and apical disc 45-52% of test length; mean
= 48%). Distinguished from other species by (i) its
depressed, rounded profile, (1) broad, relatively shallow
anterior groove (iii) central apical disc.

DESCRIPTION. Tests range from 44 to 89 mm in length and
from 38 to 68 mm in width (width = 80-88% of length; mean
= 85%, SD = 2:1%, N = 4). In outline the test is oval with a
prominent anterior sulcus and a truncated posterior (PI. 31,
figs 1, 3, 4, 6). The widest point on the test is approximately
mid-length. In profile the test is depressed, more or less
uniformly curved both in front and behind, with the tallest
point of the test anterior of centre (PI. 31, figs 2, 5). Test
height is 42-53% of length (mean = 48%, SD = 4:2%, N =
5). In anterior profile the sides are uniformly rounded.

The apical disc lies approximately mid-length, 43-52% of
test length from the anterior border (mean = 48%, SD =
2-4%, N = 15). Plating appears to be similar in arrangement
to that of H. compressus, with the anterior pair of genital
plates separated from the posterior pair by ocular plates II
and IV, which meet centrally (Fig. 78D). Madrepores extend
over genital plates 2 and 3, and over ocular plates II, III and

The pores of the anterior groove are small and closely
spaced adapically, becoming slightly more widely separated
towards the ambitus. The anterior sulcus is rather broad and
shallow, becoming more or less parallel-sided towards the
anterior border (Pl. 31, figs 1, 3, 6). The margins of the
groove are gently rounded, never crested. The width of the
sulcus is 17-22% of the test width (mean = 19%, SD = 2:0%,
N = 13) and at its deepest it is only 2:0-3-0 mm in depth. The
sulcus does not have a vertical component at the anterior, as
is seen in H. compressus, but curves uniformly towards the
ambitus.

Petals are as in H. compressus, though the anterior pair are
slightly more curved forwards than in that species (Fig. 79).
The anterior column of pore-pairs in each petal is composed
of small, rudimentary pore-pairs which become slightly larger
towards the ambitus. The posterior column in each petal is
composed of wide, conjugate pore-pairs, with the inner pore
circular and the outer pore distinctly slit-like. There are 36
pore-pairs in the posterior column of ambulacrum II petal,
and 28 in ambulacrum I petal in a 44 mm length individual,
rising to 65 and ?55 pore-pairs respectively in an 80 mm
individual. The anterior petals curve forwards to diverge at
an angle of approximately 30° to the midline, before tapering

A.B. SMITH

and turning laterally slightly at their distal end. They extend
almost to the ambitus in adapical view (Fig. 79). The poste-
rior petals are slightly shorter, extending approximately two-
thirds of the distance to the ambitus. They too are flexed, but
for most of their length they diverge at approximately 50° to
the midline. Pores below the petals are microscopic, but
remain double. Peribuccal pores are well-developed, with
four or five to a column in ambulacrum III, six or seven in
ambulacra II and IV, and four or five in ambulacra I and V.

The interambulacra become very narrow adapically (Fig.
79), but remain biserial. The plastron consists of a small,
broad, labral plate followed by five wedge-shaped plates
arranged uniserially (Fig. 79).

The peristome is oval to crescentic, slightly more than
twice as wide as long, and lies at the base of the anterior
sulcus, some 17-21% of test length from the anterior (mean
= 19%, SD = 1:8%, N = 5). The labral lip projects
downwards slightly and is visible in anterior view. The labral
plate indents the posterior of the peristome.

The periproct is oval, 65-75% as wide as tall (mean =
71%), and more or less vertical on the posterior face. It is
therefore not visible from either the apical or oral surfaces
(Pl. 31, figs 1-4). There are no subanal protuberances devel-
oped in small individuals, although small protruberances are
found on larger individuals (Pl. 32, fig. 2). The periproct
opens between interambulacral plates 5 and 10, the lower
plates being strongly curved. The periproct lies low on the
posterior surface and the distance to the base of the periproct
from the lower surface is some 16-25% of the test height
(mean = 22%).

Aboral tuberculation consists of small primary tubercles
scattered amongst dense miliaries. Larger primary tubercles
are found along the inner (interambulacral) margins of the
frontal sulcus, where they are approximately three abreast.
They are also found close to the apical disc interambulacrally
on many juveniles. These adapical tubercles are only slightly
larger than other tubercles, but form a characteristic feature
where preservation is good. There is no trace of a lateral
fasciole. Oral tubercles are slightly larger than adapical ones,
but there is no real difference in size between the tubercles of
the plastron and those of the latero-ventral regions. The
plastron tubercles are largest towards the adambulacral mar-
gins and decrease in size towards the midline. Oral ambulacra
are smooth and free of tubercles.

REMARKS. I have examined topotype material of this species
from Iran and feel secure that the Omani Mountain material
is conspecific. However, there is one small difference; the
Omani specimens in general have a slightly shallower anterior
groove at the ambitus than do the Iranian specimens. This is
not considered sufficient to merit separation, since all inter-
mediates can be found. Similarly, Hemipneustes sardanyolae
Vidal from the late Campanian of Sardanol, Spain, is slightly
flatter and more oval in outline, but is otherwise very similar.
It too is synonymized here.

The species is readily distinguished from H. compressus by
its very different, more depressed profile, shallower and very
much broader frontal sulcus and central apical disc. It also
has better developed phyllodes. It differs from H. arabicus in
its very broad and shallow frontal sulcus, and from Opisop-
neustes in the lack of aboral primary tubercles.

There is little doubt that the closest species to H. persicus is
H. delettrei (Peron & Gauthier) from the late Campanian of
North Africa. Both have a very similar broad, shallow frontal

~the

ann lem

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 229

PLATE 31

Figs 1-7 Hemipneustes persicus Cotteau & Gauthier. 1, 2, BMNH EE4084; 1, apical; 2, lateral; both x 1. Jebel Rawdah, section 3, bed 5.

3-5, BMNH EE5032; 3, apical; 4, oral; 5, lateral; all x 1. Jebel Rawdah, section 2, bed 21. 6, BMNH EE3746, apical, x 1. Jebel Rawdah,
section 2, bed 21. 7, BMNH EE3747, lateral, x 1. Jebel Rawdah, section 2, bed 21.

Figs 8-11 Hemipneustes compressus Noetling. BMNH EE3745; 8, apical; 9, oral; 10, anterior; 11, lateral; all x 1. Jebel Rawdah, section 2,
bed 21.

230

groove that expands continuously towards the anterior, a
character seen in no other Hemipneustes. H. persicus differs
from H. deletrettei primarily in being very much more
depressed in profile and more elongate, with more strongly
inflexed petals. However, the differences are not great.

Hemipneustes arabicus Ali, 1989 _ PI. 30, figs 1-4; Figs
76, 78B, 80

1989 Hemipneustes arabicus Ali: 408, figs 6 (1-3), 7.
1989 Hemipneustes persicus Cotteau & Gauthier; Ali: 408,
fig. 6 (4).

Types. The syntype series are the seven specimens and three
fragments mentioned by Ali (1989) as being housed in the
Geology Department Museum, Al Ain University, United
Arab Emirates. The lectotype, here designated, is his figured
specimen (op. cit. fig. 6 (1-3)).

MATERIAL STUDIED. Nine specimens: the following descrip-
tion is based on the four best-preserved of these, BMNH
EE4085, EE4087-88, EE5027.

OCCURRENCE. This species was first described from Jebel
Rawdah, Oman. Specimens were collected from the follow-
ing horizons at Jebel Rawdah:

Section 1: bed 4 (3).

Section 2: bed 11 (4); bed 14 (1).

Section 4: loose in scree, a little below beds 21/22 (1).

DIAGNOSIS. Test ovate and rather elongate, depressed in
side-view and in cross-section arched. Width 76-79% of
length, mean = 77%; height 45-46% of length, mean = 45%
(size range 76-89 mm). Frontal groove narrow, sharply
defined with a small but prominent rim; deeply sunken at the
anterior. Apical disc lies at or slightly behind mid-length
(distance between anterior and apical disc = 49-55% of test
length: mean = 52%). Distinguished from other species by (i)
its depressed, elongate profile, (11) long, sharp and deep
anterior groove (iii) central apical disc.

DESCRIPTION. Tests range in length from 77 to 89 mm and in
width from 60 to 69 mm (width = 76-79% of length, mean =
77%, SD = 1-:1%, N = 4). The widest point lies at or slightly
in front of mid-length. In outline the test is oval with a sharp
and narrow anterior sulcus and a rather broad posterior
truncation (PI. 30, fig. 1). In side-view the test is depressed
and gently rounded (PI. 30, fig. 4). Test height is 44-46% of
length (mean = 45%, SD = 0-8%, N = 4). The tallest point
on the test is at, or slightly in front of, the apex. In anterior
view the test is uniformly rounded, but there are small crests
on either side of the sulcus.

The apical disc is clearly seen in BMNH EE5S5027 (Fig.
78B). Genital pores are large and separated, with ocular
plates If and IV meeting centrally and separating genital
plates 2 and 3 from genital plates 1 and 4. Madrepores cover
genital plates 2 and 3 and ocular plates II, II] and IV. A
couple of pores also appear on the margin of genital plate 1.
It lies 49-55% of the test length from the anterior border
(mean = 52%, SD = 2:5%, N = 4).

Ambulacrum III is narrow and parallel-sided adapically
(Pl. 30, fig. 2). It is slightly depressed from the apex until it
approaches the anterior border, then it turns rather sharply
downwards into a deep groove, 4-6 mm in depth. The sulcus
remains narrow throughout, only 11-16% of the test width at
the anterior (mean = 13%). Pore-pairs are densely packed

A.B. SMITH

rN
a

et
et

Fig. 80 Camera lucida drawing of adapical plating of Hemipneustes
arabicus, BMNH EES027. Scale bar = 5 mm.

along its length until it turns adorally, where they become
much more spread out. The floor of the sulcus is covered in
fine tuberculation which decreases in size away from the
pore-pairs towards the perradius.

The petals are similar in form to those of other species of
Hemipneustes, the adapical pores in the anterior columns
being particularly rudimentary. The anterior petals are flexed
forward and diverge at an angle of about 15° to the frontal
groove before curving laterally near their distal end (Fig. 80).
The posterior petals diverge at about 110—120° to each other.
Adorally there are well-developed phyllodes, with five or six
pores in a column in the lateral ambulacra.

The plastron consists of a small, broad labral plate and a
series of four or five wedge-shaped plates that extend across
the full width of the plastron (Fig. 80). Posterior plates are
biserial. The peristome is oval, indented slightly by the labral
plate, and predominantly forward-facing. It lies at the base of
the anterior sulcus, some 16% of the test length from the
anterior border.

The periproct is oval and downward facing; clearly visible
from below but hidden from above (PI. 30, figs 1, 2). The test
is developed into two prominent bulges, one on either side of
the periproct. The periproct lies between interambulacral
plates 5 and 10. The base of the periproct lies between 19 and
26% of the test height above the base.

Tuberculation is fine and uniform over the apical surface,
except along the inner interambulacral margins-of the frontal
groove, where larger primary tubercles are developed. Oral
tuberculation is slightly coarser, with ambulacral zones free
of any tuberculation. No lateral fasciole is seen.

REMARKS. This species appears somewhat intermediate
between the broad grooved H. persicus and the tall, narrow-
grooved but upright H. compressus. It differs from H.
persicus in the sharpness and narrowness of its anterior

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 231

PLATE 32

Figs 1-4 Hemipneustes persicus Cotteau & Gauthier. Specimen in the Morgan Collection, Museum d’Histoire Naturelle, Paris; 1, apical; 2,
oral; 3, lateral; 4, anterior; all x 2. Arkouraj, Iran.

Figs 5-8 Hemiaster hattaensis Ali. 5, BMNH EE4060; apical, x 2. Jebel Thanais, basal 2 m of the Simsima Formation. 6-8, BMNH EE4059;
6, lateral; 7, apical; 8, oral; all x 1-5. Jebel Buhays, section 1; loose in the scree derived from the lowest 3 m of the Simsima Formation.

232

sulcus. In H. persicus the frontal groove is much broader and
lacks the interambulacral keels. It is also much less deeply
indented at the ambitus. However, like H. persicus, H.
arabicus has its apical disc at or slightly behind mid-length, in
contrast to H. pyrenaicus where the test is more quadrate and
the apical disc lies anterior of centre. Like H. compressus, H.
arabicus shows a sharp change from apical to anterior sectors
of its anterior sulcus.

Order SPATANGOIDA Claus, 1876
Family HEMIASTERIDAE Clark, 1917
Genus HEMIASTER Agassiz, in Agassiz & Desor, 1847

Pl. 32, figs 5—8; Figs
81C-E

Hemiaster hattaensis Ali, 1989

21903 Hemiaster sp. Lambert: 87, pl. 3, figs 6-8.
1989 Hemiaster (Bolbaster) hattaensis Ali: 409, fig. 6 (5-8).

Types. Eleven specimens referred to by Ali form the syntype
series since no holotype was designated. These are in the
Geology Department Museum, University of Al Ain, United
Arab Emirates.

MATERIAL STUDIED. Thirteen specimens, of which the fol-
lowing six were used for the biometric study: BMNH
EE4055, EE4057, EE4059-60, EE4064 and EE4320.

OCCURRENCE. The species is known for certain only from

the western Oman mountain. It has been found at the

following localities:

Jebel Buhays, section 1: loose in the scree, derived from the
lowest few metres of the Simsima Formation (5).

Jebel Thanais: lowest 2 metres of the Simsima Formation (1).

Jebel Rawdah, section 1: bed 6 (3).

Jebel Rawdah, section 2: loose in scree (1).

Jebel Rawdah, section 3: bed 5 (1).

Jebel Rawdah, section 4: bed 2 (1); bed 13 (1).

We ‘

Ly
Le

Fig. 81

A.B. SMITH

DIAGNOsIS. An ovate Hemiaster with cruciform petals in
which the posterior pair are about half the length of the
anterior pair. Apical disc ethmophract, with the posterior
genital plates L-shaped and separating the posterior ocular
plates; lying posterior to midlength. Ambulacrum III long,
narrow, depressed adapically but becoming flush with sur-
rounding area towards the ambitus.

DESCRIPTION. Tests oviform with a uniformly rounded ante-
rior and a slightly pointed posterior (Pl. 32, fig. 5). Test
length is 15-32 mm and test width 11-26 mm (width =
82-87% of length in larger individuals but only 75% in a
juvenile 15 mm long). The widest point lies about mid-length.
Test rather depressed in profile (Pl. 32, fig. 6), with a height
55-67% of length (mean = 64%, SD = 4-6%, N = 6). Lower
surface gently convex with a slight keel to the plastron
towards the peristome. Upper surface flat towards the poste-
rior, sloping uniformly towards the anterior (PI. 32, fig. 6).
The tallest point on the test lies just posterior to the apical
disc.

The apical disc is ethmophract (Fig. 81D) and lies 58-62%
of the test length from the anterior border in adults, but
further (73%) in the juvenile 15 mm in length. Anterior
gonopores are set closer together than the posterior pair.
Genital plates 2 and 3 are relatively small whereas genital
plates 1 and 4 are longer and L-shaped and are broadly in
contact (Fig. 81D). There are relatively few madrepores
developed. Ocular plates are small, pentagonal and project.

The anterior ambulacrum is sunken adapically but the
sulcus is lost about two-thirds of the way towards the ambitus
and there is no indentation at the ambitus (PI. 32, figs 5, 7).
The adapical sulcus is parallel-sided and very narrow. There
are 14 pore-pairs in a column in 17-18 mm individuals and
these are strongly oblique with a prominent interporal parti-
tion. The floor of the sulcus is covered in fine and dense
granulation. Pores beyond the peripetalous fasciole are single
and minute. The interambulacra on either side of the frontal
sulcus are keeled, especially in the larger specimens.

————

Camera lucida drawings of plating in Hemiaster species. A, B, Hemiaster paronai Checchia-Rispoli, BMNH EE5034: A, apical

surface (fasciole stippled); B, apical disc. C-E, Hemiaster hattaensis Ali: C, peristome and labral plate, BMNH EE4064; D, apical disc,
BMNH EE4060; E, apical surface (fasciole stippled), BMNH EE4059. Scale bars: A, E = 5 mm; B, D = 1 mm; C = 2mm.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

The petals are cruciform and sunken, the anterior pair
being twice as long as the posterior pair (PI. 32, figs 5, 7; Fig.
81E). Pore-pairs are broad and conjugate and the columns
close distally. The perradial zone is slightly narrower than the
pore-pair zones on either side. There are 12-14 pore-pairs in
a column in the posterior petals and 24-26 in the anterior
petals in individuals 15-18 mm in length.

The periproct is tear-drop shaped, pointed adapically, and
lies high on the posterior face. It is about 3 mm wide and 4
mm in height in an individual 17 mm in length. The base of
the periproct lies 55-58% of test height above the base.

The peristome is D-shaped and is 1-6 to 1-8 times as broad
as wide. It lies 24-29% of the test length from the anterior
border in 17-18 mm length individuals. It is neither invagi-
nated nor does it have a surrounding rim, as is seen in many
Hemiaster species. The plastron is broad, with a straight,
median suture (Fig. 81C). It is covered in dense orderly rows
of tubercles. The labral plate is vase-shaped and relatively
small. Surrounding the peristome there are four phyllode
pores in each column of a lateral ambulacrum and three in the
posterior ambulacrum. There are also two or three enlarged
subanal pore-pairs.

There is a well-developed peripetalous fasciole that is
without sharp angles. The remainder of the upper surface has
scattered primary tubercles and dense miliaries.

REMARKS. Ali (1989) erected this species for 11 specimens
from Jebel Rawdah. It resembles H. aquisigranensis Schluter
in the form of its petals and frontal groove, but differs from
that species in having a larger peristome and having a vertical
or slightly outward-sloping posterior. In H. aquisigranensis
the posterior is strongly retrenched, such that the periproct is
visible from the oral surface rather than the aboral surface.
H. hattopsis undoubtedly comes closest to H. punctatus
_d@Orbigny, from the late Campanian of France. However,
~ this species differs in being very much smaller, more elevated
and in having a well developed rim to the peristome.
Lambert (1903) described and figured a badly preserved
_ specimen that may be conspecific. Lambert’s specimen came
from the late Cretaceous of Fanivelona, eastern Madagascar.
_ It has a very similar overall shape, but unfortunately the apex
of the test is damaged and only the anterior petal on one side
is preserved. It is only tentatively assigned to this species.

Hemiaster paronai Checchia-Rispoli, 1921 Pl. 33, figs
1-4; Figs 81A, B

1921 Hemiaster Paronai Checchia-Rispoli: 27, pl. 8, fig. 24,
pl. 9, figs 14-18.
| 1932 Hemiaster Paronai Checchia-Rispoli;
Rispoli: 8, pl. 2, figs 1-8.

21967 Hemiaster punctatus d’Orbigny; Devries: 194, pl. 6,
. figs 34-41.

21967 Hemiaster regulusanus d@’Orbigny; Devries: 194, pl. 6,
figs 42-44.

Checchia-

MATERIAL. Seven specimens, only one of which is well-
_ preserved, BMNH EE5034.

) OCCURRENCE. Specimens were found at the following locali-
ties and horizons in the western Oman mountains:

| Jebel Huwayyah, section 1: bed 9 (1); bed 11 (1).

| Jebel Huwayyah, section 2: beds 3-5 (3).

_ Jebel Faiyah, section 1: bed 4 (1).

Jebel Rawdah, section 3: bed 9 (1).

|
|

233

Elsewhere the species has been recorded from the Maastrich-
tian of northern Libya.

DIAGNOsIs. Like H. hattaensis but more inflated and more
circular in outline, with deeper petals and deeper and wider
anterior sulcus adapically.

DESCRIPTION. The best preserved specimen is 37-1 mm in
length, 35-6 mm in width (96% of length: widest point
approximately mid-length) and 28-8 mm in height (78% of
length). In outline the test is more or less circular and in
profile the upper and lower surfaces are broad and flat with
the anterior uniformly rounded and the posterior sloping
steeply outwards (PI. 33, figs 1-4).

The apical disc is ethmophract with the posterior two
genital plates rather stout and broadly in contact (Fig. 81B).
The posterior ocular plates are separated. Gonopores are
large and occupy most of the plate, but in some individuals
(?males) may be relatively smaller. Genital plate 2 has a small
central zone of madrepores. The apical disc lies 58% of the
test length from the anterior border.

The anterior ambulacrum lies in a frontal sulcus which is
bordered by sharp interambulacral crests adapically but
which shallows and is lost towards the ambitus (PI. 33, fig. 2).
There are 22 pore-pairs in a column between the apex and the
peripetalous fasciole and these are strongly oblique. The
sulcus is rather narrow and slightly lanceolate in outline.

The petals are cruciform with the anterior pair twice the
length of the posterior pair (Fig. 81A). There are 40 pore-
pairs in a column in the anterior pair and 21 in the posterior
pair at 37 mm test length. Petals close distally and the
perradial interporal zone is narrower than either of the
bordering pore zones.

The periproct is oval, slightly pointed adapically and lies
high on the posterior side. It is just visible from above
because of the outward slope of the posterior face. It is about
70% as wide as tall and is 18% of the test height in height. Its
base lies 60% of test height above the base (PI. 33, fig. 4).

The peristome is D-shaped, twice as wide as long and lies
28% of the test length from the anterior border. It is not
rimmed. There are five phyllode pores in each column of
lateral and posterior ambulacra. There are also five subanal
pore-pairs in the two posterior interambulacra.

REMARKS. This species closely resembles H. hattaensis in the
shape of petals, apical disc structure and the form of the
anterior ambulacrum. However, it differs consistently in
shape, being both more rounded in outline and more inflated
in profile. This is not simply an attribute of size, since the
small specimens from Jebel Huwayyah (eg. BMNH EE4061,
22 mm in length) are very different in shape from similar-
sized individuals from Jebel Buhays. In addition the pore-
pairs in the frontal groove are more numerous and more
densely packed.

The specimens appear indistinguishable from _ those
described from the Maastrichtian of Libya by Checchia-
Rispoli (1921) under the name H. paronai. Very similar
material was also described by Devries (1967) from the
Maastrichtian of Turkey under the names Hemiaster puncta-
tus d’Orbigny and H. regulusanus d’Orbigny. Unlike the
specimens described here and by Devries (1967), H. puncta-
tus has a large flush peristome lacking a rim. The species is
also very similar to Hemiaster noemiae Cotteau & Gauthier,
from the late Senonian of southern Iran (Pl. 33, figs 5-8).
However, H. noemiae differs in having a shorter anterior

234 ; A.B. SMITH

PLATE 33

Figs 1-4 Hemiaster paronai Checchia-Rispoli. BMNH EES034; 1, oral; 2, apical; 3, lateral; 4, posterior; all x 2. Jebel Rawdah, section 3,
bed 9.

Figs 5-8 Hemiaster noemiae Cotteau & Gauthier. B18727, Morgan Collection, Museum d’Histoire Naturelle, Paris; 5, oral;6, apical; 7,
posterior; 8, lateral; all x 2. Senonian of Awasa, Iran.

Figs 9-11 ?Linthia sudanensis (Bather). BMNH EE4054; 9, lateral; 10, apical; 11, oral; all x 1-5. Loose near top of section at Jebel
Rawdah, section 3.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS 2B5

PLATE 34
Figs 1-3, 8 Proraster geayi Cottreau. 1-3, specimen in Museum d’Histoire Naturelle, Paris; 1, apical; 2, oral; 3, lateral; all x
Maastrichtian of Antanihody, Madagascar. 8, BMNH EE4067, apical, x 1. Jebel Huwayyah, section 1, bed 18.
| Figs 4-7 Mecaster victoris (Lambert). Jebel’ Rawdah, section 3, bed 5. 4, BMNH EE4050; apical, x 2. 5, BMNH EE4049; oral, * 25 Grills
BMNH EE404?2; 6, apical; 7, lateral; both x 2.

236

sulcus with many fewer pore-pairs, and in having a much
smaller peristome at comparable sizes.

Genus MECASTER Pomel, 1883

Mecaster victoris (Lambert, 1932) PI. 34, figs 4-7; Figs
82-84A

1932 Hemiaster Victoris Lambert: 127, pl. 4, figs 18, 19.
1967 Hemiaster sp. Devries: 194, pl. 6, figs 45—47.
1990 Periaster subsexangulatus Airaghi; Ali: 410, fig. 5 (8).

MATERIAL STUDIED. Ten reasonably complete specimens
were used in the biometric analysis, BMNH EE4035—36,
EE4038-40, EE4042, EE4045—-46, EES035-36. Another 32
specimens were collected.

OCCURRENCE. In the study area, this species was found at

the following localities and horizons:

Jebel Buhays, section 1: loose in the scree, derived from the
lowest few metres of the Simsima Formation (1).

Jebel Huwayyah, section 1: bed 14 (1).

Jebel Huwayyah, section 2: beds 3-5 (3).

Jebel Rawdah, section 3 (top of bed 5 (28): bed 8 (2); beds
9/10 (7).

The species was originally described from the Campanian
of El Kantara, Algeria. A poorly preserved specimen that
also appears identical was described from the Maastrichtian
of Cortinek, Turkey.

DESCRIPTION. Specimens range in size from 24 to 33 mm in
length. Mean test width is 98% of the length (range 93-100%,
SD = 3:2%, N = 9) and test height 76% of the length (range
72-80% , SD = 3-:1%, N = 7). The test is weakly cordiform in
outline, tapering posteriorly to a rounded point and with the
widest point in front of the mid-line (Pl. 34, figs 4-6). In
profile the test is inflated with a rounded posterior and
anterior (Pl. 34, fig. 7). The tallest point is posterior of the
apical disc.

The apical disc lies 36-40% of the test length from the
anterior border (mean = 38%, SD = 1-4%, N = 8). It is
ethmolytic with the posterior oculars separated by the
madreporite (Fig. 84A). The gonopores on each side of the
mid-line lie close together.

Although ambulacra are sunken, they are all rather shallow
and relatively narrow (PI. 34, figs 4, 6; Fig. 833A). Pore-pairs
in the anteal sulcus are small and situated along the adradial
margins. The anterior petals are long, narrow and straight-
sided. They extend most of the way to the ambitus and
diverge from one another at an angle of 130°. There are 33
pore-pairs in a column at 25 mm test length, rising to 36 at 29
mm test length. The pore-pairs are narrow, with the outer
pore in each pair slightly more elongate than the inner. The
posterior petals extend about 75% of the distance to the
ambitus and are equally narrow and parallel-sided. They are
about 85-90% of the length of the anterior petals and have 28
pore-pairs in a column at a test diameter of 25 mm, rising to
about 32 at 28 mm.

The periproct is high on the posterior and is usually just
visible from above. The base lies more or less half way up the
posterior face (mean = 50%, SD = 4:5%, N = 4). The
opening is vertically elongate, being about 1-6—1-8 times as
tall as broad, and is pointed adapically.

A.B. SMITH

The peristome is D-shaped and lies about 20% of the test
length back from the anterior border (range 18-5—21-5%, SD |
= 1:1%, N = 6). The labrum projects slightly over the |
opening. The labral plate is relatively long and narrow,
almost parallel-sided (Fig. 83B). It meets only one of the |
sternal plates.

Tuberculation is fine adapically, slightly coarser adorally. |
There is always a peripetalous fasciole, some 0-6 mm in
width. Although preservation is usually too poor to allow its
course to be traced fully, there appears to be little indentation
of the peripetalous fasciole laterally and it runs close to the |
ambitus. In one specimen (BMNH EE4050) there is a distinct |
latero-anal fasciole also, but other specimens, equally well
preserved in this region have only rudimentary traces of such
a fasciole or no fasciole at all. It is clear that the latero-anal |
fasciole is variably developed in this species.

}
REMARKS. Lambert (1932: 127) created this species on the
basis of material from the late Campanian of El Kantara, |
Tunisia. It has narrower, shallower petals than other
Mecaster species and in profile is inflated with the tallest point

lying well to the posterior. Lambert noted that there 5

traces of a latero-anal fasciolt in six out of twelve specimens, |
while another three had a distinct latero-anal fasciole. Lam-
bert separated those with a latero-anal fasciole and placed
them in his ‘Periaster Victoris’, though recognising that they
were completely intergradational with Hemiaster victoris. As
Lambert quite correctly pointed out, the presence/absence of
a latero-anal fasciole is highly variable and the various species
classified under the genus Periaster are polyphyletic in origin. |—
Ali (1989) figured a specimen of this species without .
description under the name Periaster subsexangulatus Air-
aghi. However, P. subsexangulatus has a much broader, |_
deeper anteal sulcus, is less rounded and less inflated and

most particularly, the petals are broader and less parallel- |
sided and the posterior pair are distinctly shorter than the |
anterior pair.

The plastron structure is significant in that the labral plate
is in contact with sternal plate 2b only (Fig. 83B). This is also
the situation in /raniaster and other Somaliasteridae. /rant-
aster has been placed in the Holasteroida on account of its
plastron structure, yet it has a compact apical disc in which
the posterior genital plates, but not the posterior ocular
plates, are separated by the madreporite (Kier 1972, figs
41-42). The fact that a meridoplacous plastron can be devel-
oped in at least one Mecaster species, and the similarity of
apical disc plating strongly indicates that somaliasterids are
derived from the Mecaster lineage.

ge a eS ee

Genus LINTHIA Desor, 1853
4

?Linthia sudanensis (Bather, 1904) __ PI. 33, figs 9-11;

Figs 84B, 85
21904 Hemiaster sudanensis Bather: 299, pl. 11, figs 6-13.

Family SCHIZASTERIDAE Lambert, 1905

i
MATERIAL. One specimen, BMNH EE4054.

OCCURRENCE. The specimen was found loose about 8 m
below the top of section 3 at Jebel Rawdah, western Oman.
The section here is capped by a two metre thick conglomerate
of reworked Simsima Limestone and thus the specimen could
be of late Maastrichtian age.

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

40
| = Test width
| @ Test height

22 24 26 28 30 32
Test length (mm)

gs Anterior to apical disc
@ Anterior to peristome —

Distance (mm)

Test length (mm)

| Fig. 82 Biometric data for Mecaster victoris (Lambert).

EE4049. Scale bar = 2 mm.

DESCRIPTION. The specimen is not well preserved but retains
‘sufficient features to be able to place it generically. The test is
32-5 mm in length, 30-5 mm in width (93% of length) and 24
mm in height (74% of length). It is heart-shaped in outline
‘with a squarely truncated posterior and a rather deep anteal
‘sulcus, approximately 2-5 mm deep (7% of test length). The
anteal sulcus is relatively broad and deepens gradually away
from the apex. In profile the test is rather flat above and
below and has a rounded anterior and steep posterior.

Length of petal (amb. Il) (mm)

\ raga

237
22 24 26 28 30 32 34
Test length (mm)
5.0 a a

24 26 28 30 32 34
Test length (mm)

see

| Fig. 83 Camera lucida drawings of plating in Mecaster victoris (Lambert). A, apical surface, BMNH EE4050; B, oral surface, BMNH

The apical disc is ethmolytic and gonopores are large, those
on either side almost touching (Fig. 84B). It lies well anterior
of the centre, the anterior gonopores being 38% of the test
length from the anterior border. The posterior ocular plates
are separated by the posteriorly elongated madreporite plate.
The antero-lateral ocular plates project and hardly indent the
adjacent genital plates.

The anterior sulcus has small isopores that are not crowded
together. The anterior petals are 13-5 mm in length and

Fig. 84 Camera lucida drawing of apical discs. A, Mecaster victoris
(Lambert), BMNH EE4050. B, ?Linthia sudanensis (Bather),
BMNH EE4054. Scale bar = 1 mm.

ee a

Fig. 85 Camera lucida drawing of ?Linthia sudanensis (Bather),
BMNH EE4054. A, apical; B, oral; C, lateral. Scale bar = 5 mm.

widely divergent (Fig. 85). They extend for most of the
distance to the ambitus when viewed from above. Inner and
outer pores are equally slit-like and there are 33 pore-pairs in
a column. The posterior petals are shorter, only 10 mm in
length, and reach approximately half-way towards the ambi-
tus. There are 24 pore-pairs in a column.

The periproct is just visible from above. It is 6-5 mm in
height and 4 mm in width, being pointed adapically. The base
of the periproct lies just above mid-height, some 11-3 mm
above the base of the test (52% of test height).

The plastron is asymmetric, with the median suture dis-
placed towards the right. The labral plate is short (11-5 of the
test length) and trapezoidal in outline. The peristome is
D-shaped although the labral plate does not project over the
peristome much. The peristome is set rather far back from
the anterior border, its anterior margin being 23% of the test
length from the anterior.

REMARKS. The specimen is unfortunately very badly pre-
served and cannot be identified to species level with any
certainty. However, from the general shape and petal form it
cannot be distinguished from the common Palaeocene species
Linthia sudanensis described by Bather (1904) from Sudan.

A.B. SMITH

Genus PRORASTER Lambert, 1895

Pl. 34, figs 1-3, 8 |

Proraster geayi Cottreau, 1908
1908 Proraster Geayi Cottreau: 26, pl. 4, fig. 5. |

Types. The syntypes of P. geayi are the two specimens
described from Marohita, eastern Madagascar by Cottreau
(1908) stated to be in the Museum d’Histoire Naturelle,
Paris.

MATERIAL STUDIED. Six specimens, BMNH EE4065-69,
EE5037, none of which are particularly well-preserved.

occurs as juveniles in beds 3—5 at Jebel Huwayyah, section 2
(2). Large individuals occur in bed 18 at the top of section 1,
Jebel Huwayyah (7), and in bed 9 at Jebel Rawdah, section 1 |
(2). It is also found in the Maastrichtian of Madagascar.

|
OCCURRENCE. In the western Oman Mountains this species |
\

DIAGNOsIs. A Proraster with a very deep anteal sulcus that is
closed or almost closed distally by convergence of the lateral —
walls.

DESCRIPTION. No test is well-preserved, the best specimens
being BMNH EE4067 and EE4068. The former is a large
individual 71 mm in length, 56 mm in width and about 27 mm
in height, whereas the latter is a much smaller individual 27
mm in length, 24 mm in width and 15-6 mm in height. The
test is oval in outline with its widest point slightly posterior of
midline (Pl. 34, fig. 8). In profile the test is depressed and

wedge-shaped, sloping gradually towards the anterior. The |
posterior is almost vertical and the tallest point on the test lies |
close to the posterior.

The apical disc lies well towards the posterior, 66-69% of
test length from the anterior border. Plating is not seen in any
specimen.

The anterior ambulacrum is extremely deep and at the) ~
ambitus is about 10% of the test length in depth. The walls
are concave with the adjacent interambulacra curving over | |
the groove. Towards the anterior the two sides almost touch |
so as to roof over the sulcus. The groove thus appears
pinched shut close to the anterior (PI. 34, figs 1, 3, 8).

Petals are sunken. The anterior pair curve forward and run) -
parallel with the anteal sulcus for about two thirds of its) —
length. The posterior petals are very much shorter (about one
third of the length of the anterior petals) and diverge strongly
at about 120°. There is a well developed peripetalous fasciole
developed at the base of the petals, which is presumably |
continuous, but is not seen other than in small patches
because of poor preservation. There is no evidence of a} |
latero-anal fasciole. 3

The peristome is positioned far forward, lying at the base) »
of the anteal sulcus, and faces anteriorly. The periproct is) ~
large and situated high on the posterior face. |

REMARKS. Although the Omani specimens to hand are not
well-preserved they show sufficient detail to be unambiguously |,
assigned to this species. P. geayi was first erected for large| .
individuals from the Maastrichtian of Madagascar by Lambert
(1908). Lambert (1905) had previously erected the genus Pro-
raster for Schizaster atavus (Arnaud) from the Campanian of
Charante, France and two Iranian species Opissaster morgani
(wrongly cited as O. Douvillei) and O. centrosus Cotteau &| ~
Gauthier (1895). The genus was erected to encompass Schizaster-
like forms lacking a latero-anal fasciole.

The species of Cotteau & Gauthier are unfortunately based

LATE CAMPANIAN-MAASTRICHTIAN ECHINOIDS

on small individuals and are thus not directly comparable
with P. geayi. Differences between P. morgani and P.
centrosus seem slight and restudy may prove them synony-
mous. Because they are small, there is the possibility that
they represent juveniles of P. geayi. However, this seems
unlikely since they have much wider and less pinched anterior
sulci than very slightly larger individuals of P. geayi found in
the lower beds at Jebel Huwayyah. This pinching of the
frontal groove, whereby the two sides converge and almost
touch near the anterior border, becomes very much more
pronounced in the larger individuals higher in the section.
This character serves to distinguish P. geayi from both P.
atavus, and an undescribed ?Campanian species from Nafun,
Oman. Cotteau & Gauthier’s species also have a much less
well developed anterior notch and their peristome is standard
in orientation rather than being subvertical and directed
towards the anterior.

It is noteworthy that at Jebel Huwayyah only small indi-
viduals are found in the lower levels, in the Loftusia facies,
while large individuals are found near the top of the section in
carbonate marls representing shelf basinal facies. This could
be because the two samples represent different species with
consistent morphological differences, or because the smaller
individuals are juveniles of the larger, but inhabit a different
biotope. Without considerably more and better preserved
material it is impossible to determine which is correct. For the
present I treat the two forms as juveniles and adults of the
same species, partially because the small forms are much
closer in appearance to the large forms than they are to, for
example, P. morgani.

REFERENCES

Agassiz, A. 1863. List of the echinoderms sent to different institutions in
exchange for other specimens. Bulletin of the Museum of Comparative
Zoology, Harvard, 1 (2): 17-28.

Agassiz, L. 1836. Prodrome d’une monographie des Radiaires. Mémoires de la
Société des Sciences naturelles, Neuchatel, 1: 168-199.

— 1838. Monographie d’Echinodermes vivans et fossils. Premiére monogra-
phie: Des Salénies. 32 pp., 5 pls. Neuchatel.

— 1840. Catalogus systematicus ectyporum echinodermatum fossilium musei
Neocomiensis. Petitpierre, Neuchatel, 20 pp.

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Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):241-250

Issued 30 November 1995

Maastrichtian ammonites from the United
Arab Emirates-Oman border region

W.J. KENNEDY

Department of Earth Sciences, University of Oxford, Parks Road, Oxford OX1 3PR.

INTRODUCTION

The ammonites described are from the late Cretaceous
Qahlah and Simsima Formations of the United Arab
Emirates-Oman border region and come from two collec-
tions. Some of the specimens were collected by Dr. P.W.
Skelton (Open University) in 1990 and are housed in Oxford
University Museum (OUM prefixes). The remainder are
housed in the Natural History Museum, London (BMNH
prefixes) and were mostly collected by A.B. Smith and N.J.
Morris during fieldwork in 1991 and 1992. A few specimens
were collected by amateur enthusiasts and subsequently
donated to the Natural History Museum.

Wherever possible, ammonites have been tied down to
specific levels within measured sections. Details of the locali-
ties of these sections, and measured lithological logs are given
in a preceding section (Smith ef al.).

SYSTEMATIC DESCRIPTIONS

Order AMMONOIDEA Zittel, 1884
Suborder AMMONITINA Hyatt, 1889
Superfamily DESMOCERATACEAE Zittel, 1895
Family DESMOCERATIDAE Zittel, 1895
Subfamily DESMOCERATINAE Zittel, 1895
Genus DESMOPHYLLITES Spath, 1929

(= Schliiteria de Grossouvre, 1894: 126 (non Fritsch in
Fritsch & Kafka, 1887: 33); Schliitericeras Collignon, 1938: 92
(non Hyatt, 1903: 110))

TYPE SPECIES. Desmoceras larteti Seunes, 1892: 19, pl. 12 (3),
fig. 2; pl. 13 (4), figs 2, 3, by subsequent designation by Spath
1921: 46, as type species of Schliiteria de Grossourvre, of

’ which Desmophyllites is the replacement name.

_ Desmophyllites diphylloides (Forbes, 1846) Plate 1, figs
il, 2

1846 Ammonites diphylloides Forbes: 105, pl. 8, fig. 8.
1992 Desmophyllites diphylloides (Forbes); Kennedy &

| Henderson: 405, pl. 6, figs 1-9; pl. 16, figs 1-3, 7-8; pl.
17, figs 4~7; fig. 3F (with full synonymy).

19834 Desmophyllites diphylloides (Forbes); Kennedy &

Cobban: 120, pl. 1, figs 1-8; text-fig. 5c.

TyPEs. Lectotype, by the subsequent designation of Matsu-

moto & Obata (1955: 122) is BMNH C22682, the original of
Forbes 1846: pl. 8, fig. 8; paralectotypes are BMNH
C22683-85, all from the Upper Maastrichtian Valudavur
Formation of Pondicherry, southern India.

DESCRIPTION. BMNH C9399? consists of 270° of body cham-
ber and the nucleus of an individual with an estimated
original diameter of 62 mm. Coiling is very involute, with a
tiny, pitlike umbilicus, the umbilical wall subvertical and
narrowly rounded. The whorl section is compressed, with
whorl breadth to height ratio 0-71, the greatest breadth
around mid-flank, the flanks feebly convex and subparallel,
and the ventrolateral shoulders and venter evenly rounded.
The surface of replaced shell and internal mould are smooth,
but for a single constriction approximately 70° from the
aperture. This is narrow, shallow and markedly prorsiradiate,
straight on the inner flank, feebly convex on the outer flank
and concave on the outermost flank. Sutures not seen.

DISCUSSION. The types of Desmophyllites diphylloides are all
rather small (Kennedy & Henderson 1992, pl. 6, figs 1-9; pl.
16, figs 1-3, 7, 8), but Henderson and McNamara (1985: 54,
pl. 4, figs 1-4) described larger specimens comparable to the
present individual from the Upper Maastrichtian of Western
Australia. These show constrictions that are flexuous on the
flanks, as with the present material. The other Maastrichtian
Desmophyllites is D. larteti (Seunes, 1892) (p. 19, pl. 12 (3),
fig. 2; pl. 13 (4), figs 2, 3; see Hancock & Kennedy 1993: 154,
pl. 2, figs 1-3, 10, 11, 14; pl. 3, figs 1, 4, 5), which is a larger,
much more compressed species with a narrower, arched
venter and much more prominent and markedly fiexuous
constrictions.

OCCURRENCE. The Oman specimen is from bed 21, Jebel
Rawdah, section 2. Elsewhere, the species ranges from
Lower Santonian to Upper Maastrichtian. There are records
from southern India, Western Australia, Japan; Alaska,
British Columbia, California, and Arkansas in the U.S.A.;
Argentina, Angola; Pondoland and Zululand (South Africa),
Tunisia, and southern France (Corbieres).

Family KOSSMATICERATIDAE Spath, 1922
Subfamily KOSSMATICERATINAE Spath, 1922
Genus BRAHMAITES Kossmat, 1897

TYPE SPECIES. Ammonites Brahma Forbes, 1846: 100, pl. 8,
fig. 1, by original designation.

PLATE 1

Figs 1,2 Desmophyllites diphylloides (Forbes, 1846). BMNH C93992, Simsima Formation, Jebel Rawdah, section 2, bed 21.
Figs 3-7 Pachydiscus (Pachydiscus) neubergicus neubergicus (Hauer, 1858). 3, 4, OUM KY 1991, from an unknown level in the Simsima

Formation, southern embayment of Jebel Rawdah. 5, 6, OUM KY 1990, from an unknown level in the Simsima Formation, southern |
embayment of Jebel Rawdah. 7, OUM KY 1995, from the basal part of the Simsima Formation, Jebel Huwayyah, close to where the road |
cuts the north-western part of the outcrop.

All figures are < 1. ,

MAASTRICHTIAN AMMONITES

Subgenus ANABRAHMAITES Yabe & Shimizu, 1924
(= Subbrahmaites Yabe & Shimizu, 1924: 75).

TYPE SPECIES. Ammonites vishnu Forbes, 1846: 100, pl. 7,
fig. 9, by original designation.

Brahmaites (Anabrahmaites) vishnu (Forbes, 1846) Pl.
2, figs 9-14

1846 Ammonites vishnu Forbes: 100, pl. 7, fig. 9.

1992 Brahmaites (Anabrahmaites) vishnu (Forbes);
Kennedy & Henderson: 418, pl. 6, figs 25, 26; pl. 9,
figs 5—7, 17-20; pl. 10, fig. 5; pl. 17, figs 8, 10-11 (with
full synonymy)

Brahmaites (Anabrahmaites} vishnu (Forbes); Ken-
nedy & Hancock: 582, pl. 1, figs 5, 6.

TYPES. Lectotype, by the subsequent designation of
Kennedy & Henderson (1992: 418), is BMNH C51026, the
original of Forbes 1846, pl. 7, fig. 9; BMNH C51027 is a
paralectotype. Both are from the Upper Maastrichtian Valu-
davur Formation of Pondicherry, southern India.

DESCRIPTION. BMNH C93892 (Pl. 2, figs 11-14) consists of
200° of adult phragmocone and body chamber with a maxi-
mum preserved diameter of 83 mm, and part of the septate
inner whorls, 48 mm diameter. Coiling is very evolute, the
shallow umbilicus comprises 55% of diameter, with a broadly
rounded umbilical wall and shoulder. The whorl section is
depressed reniform, with greatest breadth just outside the
umbilical shoulder, and the whorl breadth to height ratio is
1-1. The inner whorls (PI. 2, figs 13, 14) are badly preserved,
but traces of delicate crowded ribs are present on one flank.
Both flanks have well-developed, prorsiradiate, deep, narrow
constrictions, 2 per half whorl, flanked by a strong adapical
collar-rib with feeble umbilical bulla, and a much weaker
adapertural rib, the ribs extending across the venter. The
outer whorl of the specimen is worn, but shows similar
constrictions and collar ribs, with traces of coarse prorsiradi-
ate flank ribs between constrictions. BMNH C93891 is a
much larger body chamber fragment (PI. 2, figs 9, 10), with
maximum preserved whorl height 27-7 mm and whorl breadth
to height ratio 1-0. There is a single strong constriction,
flanked by a bullate adapical collar rib that persists over the
venter, where it is strengthened into an incipient bulla (the
specimen is worn). There is a much weaker adapertural
collar-rib, and blunt non-bullate ribs, straight and prorsiradi-
ate on the flanks, weakened and feebly convex across the
venter, cover the remainder of the fragment. Sutures not
seen.

1993

DISCUSSION. These fragments are referred to Anabrahmaites
rather than Brahmaites sensu stricto on the basis of the
absence of bullae on intermediate ribs and the development
of an incipient siphonal bulla. The inner whorls of BMNH
C93892 differ in no respects from those of the paralectotype
of B. (A.) vishnu figured by Kennedy & Henderson (1992, pl.
10, fig. 5); the outer whorls of this specimen and BMNH

/ €93891 also find a match in the Pondicherry material (see

also Stoliczka 1865, pl. 79, fig 5).

OCCURRENCE. The Oman specimens are from bed 3 or 4,
Jebel Rawdah, section 1. The types are from the Upper
Maastrichtian of southern India; the holotype of Puzosia
haugi Seunes, 1892, a synonym, is from the Upper Maastrich-
tian fresvillensis Zone of southern France. The species also

243

occurs in the Maastrichtian of northern Spain and Armenia.

Family PACHYDISCIDAE Spath, 1922
Genus PACHYDISCUS Zittel, 1884
Subgenus PACHYDISCUS Zittel, 1884

TYPE SPECIES. Ammonites neubergicus Hauer, 1858: 12, pl.
2, figs 1-3; pl. 3, figs 1, 2, by the subsequent designation of de
Grossouvre 1894: 177.

Pachydiscus (Pachydiscus) dossantoi (Maury, 1930) PI.
3: BIA fhigsrie 2

Parapachydiscus dossantoi Maury: 136, pl. 16, fig. 1;
pl. 17, figs 1, 2.

1930

21944 Parapachydiscus sp. Olsson: 107, pl. 16, fig. 1.
1985 Pachydiscus (Pachydiscus) dossantoi (Maury, 1930);
Zaborski: 20, figs 17, 18, 20.
Types. Maury (1930: 136, pl. 16, fig. 1; pl. 17, figs 1, 2) based

this species on a number of specimens, referring to an
individual 190 mm in diameter as the type. All are from the
Maastrichtian on the right bank of Rio Gramame, Fazenda
do Congo, Parahyba do Norte, Brazil.

DESCRIPTION. Large, septate to whorl heights of up to 110
mm. Evolute, with 50% of previous whorl covered; umbilicus
broad, shallow, with flattened, outward-inclined umbilical
wall and broadly rounded umbilical shoulder. Whorl section
compressed, with whorl breadth to height ratio of 0-8 in the
best-preserved specimen; greatest breadth low on broadly
convex flanks; outer flanks convergent; ventrolateral shoul-
ders and venter arched. Coarse, distant ribs arise at blunt
umbilical bullae, are straight and prorsiradiate across the
flank, alternating regularly with shorter ribs that arise on the
outer flank. Ribs sweep forwards across the ventrolateral
shoulders and are weakened and broadly convex across the
venter. Ornament of this type extends to the end of the
phragmocone in BMNH C93895-96 and onto the body cham-
ber in BMNH C93894. Suture poorly preserved (Pl. 3);
intricately and deeply subdivided, as is typical for the genus.

DISCUSSION. Compressed whorl section plus persistence of
coarse, alternately long and short ribs to a large size show
these specimens to belong to P. (P.) dossantoi. Of other
Maastrichtian species, P. (P.) neubergicus (Hauer, 1858) (see
revision in Kennedy & Summesberger 1986), P. (P.) gollevil-
lensis (d’Orbigny, 1850) (see revision in Kennedy 1986) and
P. (P.) egertoni (Forbes, 1846) (see revision in Kennedy &
Henderson 1992) are also compressed, but all are more
delicately ribbed, are mature at smaller diameters, and have
adult growth stages characterized by loss of outer flank and
ventral ornament. P. (P.) jacquoti Seunes, 1890 (see revision
in Kennedy 1986) has a depressed whorl section, distant ribs
and effacement of outer flank and ventral ornament on the
phragmocone.

OCCURRENCE. Most of the Oman specimens come from the
lower Loftusia-rich beds at Jebel Huwayyah. BMNH C93894
comes from bed 10 or 11, section 1, Jebel Huwayyah, while
C93895 comes from bed 10. OUM K1998 (= Skelton 84/32:-2)
is also from Jebel Huwayyah, section 1, from an unspecified
level. One specimen, BMNH C93896, comes from a loose
block derived from the basal bed of the Simsima Formation at
Jebel Buhays, section 1. The type occurrence is of Maastrich-

244

tian age, but is difficult to place more precisely within the
stage. Kennedy (1986: 44) regarded Pachydiscus sumneri
Maury, 1930 (p. 155, pl. 13, figs 1, 2), Parapachydiscus
poseidon Maury, 1930 (p. 155, pl. 15) and Canadoceras
riogramense Maury, 1930 (p. 169, pl. 21, fig. 2), which are
said to occur in the same unit as P. (P.) dossantoi, as possible
synonyms of the Upper Maastrichtian Anapachydiscus fresvil-
lensis (Seunes, 1890). The original figures are so poor,
however, and the relative position of species within Maury’s
‘grey limestone’ is unknown, so that only the possibility of an
Upper Maastrichtian date can be considered. P. (P.) dossan-
toi from Nigeria (Zaborski 1983, 1985) comes from the
Nkoporo Shale, and was regarded as ‘probably Lower Maas-
trichtian’ (no definition of the Lower/Upper Maastrichtian
boundary was given). It co-occurs with Gaudryceras beantaly-
ense Collignon, 1956, Baculites sp. and Sphenodiscus lobatus
costatus Zaborski, 1982. On this evidence it can be dated no
more precisely than Maastrichtian.

Pachydiscus (Pachydiscus) neubergicus neubergicus
(Hauer, 1858) Plate 1, figs 3-7

1858 Ammonites neubergicus Hauer: 12 (pars), pl. 2, figs
1-3 (non pl. 3, figs 1, 2).

Pachydiscus (Pachydiscus) neubergicus neubergicus
(Hauer, 1858); Hancock & Kennedy: 158, pl. 3, figs 6,
7; pl. 9, figs 5-8; pl. 12, figs 7-9; pl. 13, figs S—7 (with
synonymy).

1993

Types. Lectotype, by the subsequent designation of de Gros-
souvre 1894: 209, is no 1858.01.6 in the collections of the
Geologisches Bundesanstalt, Vienna; three paralectotypes
bear the same number, and all are from the Lower Maastrich-
tian of Neuberg, Steiermark, Austria.

DESCRIPTION. The best-preserved specimen is OUM KY
1990 (PI. 1, figs 5, 6), an internal mould of a phragmocone 93
mm in diameter. Coiling is fairly involute, the umbilicus
small, with a flattened, subvertical low wall and narrowly
rounded umbilical shoulder. The whorl section is com-
pressed, with a whorl breadth to height ratio of 0-74, the
greatest breadth below mid-flank, inner flanks feebly convex,
outer flanks flattened and convergent, ventrolateral shoul-
ders broadly rounded, and the venter only feebly convex.
There are an estimated 14 umbilical bullae per whorl. These
give rise to single ribs, sometimes feebly concave on the
innermost flank. They are prorsiradiate and weak across the
flanks, where long and short intercalated ribs are inserted, so
that there are many more coarse concave ribs at the ventro-
lateral shoulder, although the number per whorl cannot be
determined. OUM KY1994 is a worn but conspecific frag-
ment of phragmocone with a maximum preserved whorl

PLATE 2

W.J. KENNEDY

height of 36 mm and whorl breadth to height ratio of
approximately 0-89.

OUM K1991-93 are fragments of body chamber (PI. 1, figs
3, 4), with a maximum preserved whorl height of 51-5 mm
and whorl breadth to height ratio of 0-75. At the adapical end
of the fragment strong bullate primary ribs at the umbilical
shoulder correspond to 3-4 times as many ribs at the ventro-
lateral shoulder. On the adapical part of the fragment the
secondary ribs are lost, and the ornament is of distant
primaries that efface across the flanks. OUM K1995 (PI. 1,
fig. 7) is a very worn individual 150 mm in diameter, probably
adult, with a phragmocone diameter of 110 mm and whorl
breadth to height ratio of 0-78. There appear to be 14-16
primary ribs per whorl, and more numerous primary plus
secondary ribs at the ventrolateral shoulder. Sutures not
seen.

DISCUSSION. Coiling, whorl proportions and ribbing show
these specimens to belong to the Pachydiscus (P.) neuber-
gicus neubergicus (Hauer, 1858) — P. (P.) gollevillensis
(d’Orbigny, 1850) group. Topotypes of the former generally
have 14-17 umbilical bullae and 58-60 ventral ribs per whorl
(Kennedy & Summesberger 1986), the latter 9-11 umbilical
bullae and approximately 80 ventral ribs (Kennedy, 1986),
suggesting the present material belongs to the former. Nuclei
of P. (P.) neubergicus neubergicus and P. (P.) neubergicus
dissitus Henderson & McNamara, 1985, are identical, but the
latter has a mature ornament with many ventral ribs (Hender-
son & McNamara 1985, pl. 7, fig. 7), not seen in the present
material (Pl. 1, figs 3, 4), which are thus referred to the
nominate subspecies.

OCCURRENCE. OUM KY1990, 1991-93 (1 specimen) and
KY 1994 were collected in the southern embayment of Jebel
Rawdah. Their position in the succession is unknown.
KY1995 comes from the lower part of the Simsima formation
in the north-western part of Jebel Huwayyah, close to where
the road cuts the outcrop. Elsewhere the species first appears
low in the Lower Maastrichtian, and is best known from the
Lower Maastrichtian of Austria, Poland, Ukraine, Armenia,
Russia, SW France, northern Spain, Nigeria, Brazil and
Zululand (South Africa). It occurs in the lower Upper
Maastrichtian of Denmark, and the Upper Maastrichtian of
southern India.

Indeterminate pachydiscid

Discussion. BMNH C93987 is a fragmentary pachydiscid,
still septate at a whorl height of 150 mm. It bears distant
primary ribs, but is indeterminate even as to genus.

OCCURRENCE. The specimen comes from the gritty calcare-
ous bed (bed 9), of the Qahlah Formation, immediately

Figs 1-3, 15, 16 Lewyites ambindense (Collignon, 1971). 13, BMNH C93890; from the Loftusia-rich beds, Qahlah Formation, Jebel
Huwayyah, section 1, bed 10 or 11. 15, 16, OUM KY1996; from the basal part of the Simsima Formation, north-western Jebel Huwayyah,

close to where the road cuts the outcrop.

Figs 4-8 Nostoceras (Nostoceras) major Kennedy and Cobban, 1993. 4-6, BMNH C€93994, from beds 10 or 11, Qahlah Formation, Jebel
Huwayyah, section 1; 7, 8, BMNH C93993, Basal Simsima Formation, bed 9, Jebel Bu Milh, section 2.
Figs 9-14 Brahmaites (Anabrahmaites) vishnu (Forbes, 1846). 9, 10, BMNH C93891; 11-14, BMNH C93892. Both from the Simsima

Formation, bed 3 or 4, Jebel Rawdah, section 1.

Figs 17-19 Libycoceras? sp., BMNH C93887, from the conglomeratic basal bed of the Simsima Formation, Jebel Buhays, section 2.
Fig. 20 Nostoceras (Nostoceras) sp., BMNH C93888, from bed 6, Jebel Huwayyah, section 2.

All figures are x 1

MAASTRICHTIAN AMMONITES

246 W.J. KENNEDY

PLATE 3
Pachydiscus (Pachydiscus) dossantoi (Maury, 1930). BMNH C93896, from the basal bed of the Simsima Formation, Jebel Buhays, section 1; _
slightly reduced.

MAASTRICHTIAN AMMONITES

below the Loftusia-rich beds at Jebel Huwayyah, section 1.

Superfamily ACANTHOCERATACEAE de Grossouvre,
1894
Family SPHENODISCIDAE Hyatt, 1900
(= Libycoceratidae Zaborski, 1982: 306)
Genus LIBYCOCERAS Hyatt, 1900

TYPE SPECIES. Sphenodiscus ismaelis Zittel, 1895, p. 451,
text-fig. 631, by original designation.

Libycoceras ? sp. Plate 2, figs 17-19

DESCRIPTION. BMNH C93887 is a very corroded internal
mould of a small adult with 240° of body chamber preserved
and a maximum diameter of 80 mm. The phragmocone is
oxycone, with a whorl breadth to height ratio of 0:4. The
body chamber develops subparallel flanks, and the venter
rounds progressively; the whorl breadth to height ratio is 0-39
at the aperture. No ornament is preserved on the heavily
corroded surface, other than a faint trace of low ribs on the
outer flank. The suture (Text-fig. 1) shows a well-developed
adventitious lobe in the first lateral saddle, the ventral
saddles are feebly indented, and the umbilical saddle is
entire.

DISCUSSION. This poorly preserved specimen is referred to
Libycoceras? rather than Sphenodiscus on the basis of its
sutural characteristics; it is specifically indeterminate.

OCCURRENCE. BMNH (C93887 is from the basal conglomer-
atic bed of the Simsima Formation at Jebel Buhays, section 2.
Species of Libycoceras first occur in the Upper Campanian
and may range into the Lower Maastrichtian (Zaborski
1982).

Suborder ANCYLOCERATINA Wiedmann, 1966
Superfamily TURRILITACEAE Gill, 1871
Family DIPLOMOCERATIDAE Spath, 1926
(= Neocrioceratinae Spath, 1953)
Subfamily DIPLOMOCERATINAE Spath, 1926
(= Scalaritinae Ward, 1976: 455)

Genus GLYPTOXOCERAS Spath, 1925
(= Neohamites Brunnschweiler, 1966)

| TYPE SPECIES. Hamites rugatus Forbes, 1846: 116, pl. 11, fig.
6, by original designation (Spath 1925: 30, as Hamites (Aniso-
ceras) rugatus (Forbes) Kossmat).

A 10mm ;

Fig. 1 Idealised suture of Libycoceras? sp., based on BMNH
C93887, a badly corroded specimen.

247

Glyptoxoceras sp.

DESCRIPTION AND DISCUSSION. BMNH C93889 is a straight
fragment with a whorl height of 6-5 mm, the whorl section is
compressed oval, with strong even prorsiradiate flank ribs,
transverse on the venter and weakened on the dorsum. It is
specifically indeterminate, but ribbing and coiling suggest
that it is a Glyptoxoceras, a genus recorded from Lower
Santonian to Upper Maastrichtian.

OCCURRENCE. BMNH C93889 is from the basal 60 cm shell
bed immediately overlying unweathered ophiolite (bed 1) at
Jebel Aqabah.

Genus LEWYITES Matsumoto & Miyauchi, 1984

TYPE SPECIES. Idiohamites (?) oronensis Lewy, 1969: 127, pl.
3, figs 10, 11, by original designation.

Lewyites ambindense (Collignon, 1971) Plate 2, figs 13,
5; 16

1971 Neancyloceras ambindense Collignon: 11, pl. 644, fig.
2380.

TyPe. Holotype, by monotypy, is the original of Collignon
1971, pl. 644, fig. 2380, from the Maastrichtian of locality 504
of Collignon, Mont Ambinda-Mikoboko (Manera), Mada-
gascar.

DESCRIPTION. BMNH C93890 is a 45 mm long fragment of a
straight shaft showing slight curvature at the adapical end.
The fragment is wholly septate, with traces of recrystallized
shell. The whorl section is compressed oval, with a whorl
breadth to height ratio of 0-9 and maximum preserved whorl
height of 20 mm. There are 11 ribs in a distance equal to the
whorl height. They are narrower than the interspaces, sharp,
transverse to feebly convex on the dorsum, sweeping for-
wards and feebly convex on the dorsolateral margin, mark-
edly prorsiradiate and strengthening across the flank, and
transverse on venter. Alternate ribs bear small ventral clavi;
occasionaly a second rib is feebly linked to a clavus. Much
larger is OUM KY1996, a body chamber fragment with a
maximum preserved whorl height of 29 mm. Ornament is as
in the smaller specimen, but for the marked effacement of
ribs on the dorsum of the internal mould. Sutures not seen.

Discussion. I was unable to trace the holotype of Neancylo-
ceras ambindense during a recent examination of the Colli-
gnon Collection, housed in the Université de Bourgogne,
Dijon. The ribs of the Oman material are more markedly
prorsiradiate than in the holotype, with fewer non-
tuberculate ribs, but they are otherwise similar. Reference to
Lewyites is based on a comparison with topotypes of the
type species in the Oxford University Museum (OUM
KY2021—25), and large fragments from New Jersey figured
by Cobban (1974, pl. 10, figs 22-35) and Kennedy & Cobban
(1993b, figs 5.1-5.18, 5.22-5.26; 7.19, 7.20; 9.4, 9.7).

OCCURRENCE. BMNH_ C93890 is from bed 10, Jebel
Huwayyah, section 1. OUM K1996 (Skelton 84/16-2) comes
from the basal part of the Simsima Formation of the north-
western part of Jebel Huwayyah, close to where the section is
cut by the road. The holotype is from the so-called Lower
Maastrichtian Zone a Pachydiscus gollevillensis et P. neuber-
gicus of Collignon, but associated ammonites from the type

248

PLATE 4

Pachydiscus (Pachydiscus) dossantoi (Maury, 1930). BMNH C93895 from the Qahlah Formation, bed 10, Jebel Huwayyah, section 1, x 0-75. |

locality in the Collignon Collection indicate Upper Maastrich-
tian only.

Family NOSTOCERATIDAE Hyatt, 1894
(= Jouaniceratidae Wright, 1952: 218; Bostrychoceratinae
Spath, 1953: 16;
Emperoceratinae Spath, 1953: 17; Hyphantoceratinae Spath,
1953: 16)
Genus NOSTOCERAS Hyatt, 1894
Subgenus NOSTOCERAS Hyatt, 1894

TYPE SPECIES. Nostoceras stantoni Hyatt, 1894: 569 (=
Ammonites approximans Conrad, 1855: 266), by original
designation.

W.J. KENNEDY

Nostoceras (Nostoceras) major Kennedy & Cobban,
1993 Plate 2, figs 4-8)

1993c Nostoceras (Nostoceras) major Kennedy & Cobban: |
6-1, fig. 4. |

Type. Holotype in Texas Memorial Museum Collections no}
77981, from the Upper Maastrichtian Corsicana Formation |
1-2 km (0-75 miles) SE of New Sweden, Travis County, |
Texas.

DESCRIPTION. There are 2 fragments, BMNH C93993, with a
whorl height of 13 mm, and BMNH C€93994, with a whorl |
height of 39-5 mm. Both are derived from helices with the
whorls in tight contact, there being a shallow concave
|
|

MAASTRICHTIAN AMMONITES

impressed zone on the upper whorl face; the outer and lower
whorl faces are broadly convex, the inner whorl face is
flattened. Ornament is effaced on the impressed zone of the
upper whorl face, but wiry narrow ribs strengthen, sweep
back and are concave across the juncture of upper and outer
whorl faces and are markedly prorsiradiate and narrower
than the interspaces on the outer whorl face, sweeping
backwards and feebly concave across the juncture of outer
and lower whorl faces, straight and prorsiradiate on the lower
whorl face and convex and effaced on the inner. All ribs are
single on the larger fragment, but occasionally join in pairs at
the juncture of outer and lower whorl faces in the smaller
fragment. Sutures not seen.

DISCUSSION. The simple coiling and ornament of these frag-
ments occurs in nostoceratids from the Turonian onwards.
Given the Maastrichtian age of the specimens, reference to
Nostoceras (Nostoceras) is indicated. The fragments differ in
no significant respect from the holotype of N. (N.) major,
other than their coiling direction, absence of flared ribs and
constrictions, possibly reflecting no more than the short
lengths preserved, there being only 2-3 constrictions and
associated flared ribs per whorl in the type.

OCCURRENCE. BMNH C€93993 comes from bed 9, Jebel Bu
Milh, section 2: C93994 comes from bed 10 or 11, Jebel
Huwayyah, section 1. The holotype is from the Upper
Maastrichtian of Texas.

Nostoceras (Nostoceras) sp. Plate 2, fig. 20

DESCRIPTION AND DISCUSSION. BMNH C93888 is an poorly
preserved U-shaped body chamber with a maximum pre-
served whorl height of 21 mm. Ornament is of coarse single
ribs with traces of ventral tubercles on at least some ribs. The
specimen is specifically indeterminate, but recalls the Nosto-
ceras (N.) hyatti group, of the uppermost Campanian Lower
Maastrichtian (Kennedy & Cobban 1993b).

OCCURRENCE. The specimen comes from bed 6, Jebel
Huwayyah, section 2.

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Spath, L.F. 1921. On Upper Cretaceous Ammonoidea from Pondoland.
Annals of the Durban Museum, 3: 39-56, pls 5-9.

1922. On the Senonian ammonite fauna of Pondoland. Transactions of the
Royal Society of South Africa, 10: 113-147, pls 5-9.

— 1925. On Senonian Ammonoidea from Jamaica. Geological Magazine, 62:
28-32, | pl.

— 1926. On new ammonites from the English Chalk. Geological Magazine,
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— 1929. Corrections of cephalopod nomenclature. Naturalist, 871: 269-271.

— 1953. The Upper Cretaceous Cephalopod fauna of Grahamland. Scientific
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Stoliczka, F. 1865. The fossil Cephalopoda of the Cretaceous rocks of southern

250

India. Palaeontologia Indica, Calcutta, Series 3, part 9: 143-154, pls 71-80.

Ward, P.D. 1976. Upper Cretaceous Ammonites (Santonian-Campanian) from
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W.J. KENNEDY

Zaborski, P.M.P. 1982. Campanian and Maastrichtian sphenodiscid ammonites
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Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):251-255

Issued 30 November 1995

Maastrichtian nautiloids from the United
Arab Emirates-Oman border region

NOEL J. MORRIS

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD

Synopsis. The two nautiloids Deltoidonautilus salisfilius sp. nov. and Cimomia aff. sowerbyana (d’Orbigny) are
described from the Maastrichtian Simsima Formation of the United Arab Emirates and Oman.

INTRODUCTION

Cephalopods form only a tiny percentage of the macrofossils
collected from the late Cretaceous of the Emirates-Oman
border area. They consist only of ammonites and nautiloids
and, in spite of the small numbers, it is apparent that the
distribution of the two groups is rather different and may
reflect their partial ecological separation.

Two distinct species of nautiloids were collected from a
number of the Maastrichtian localities on the Emirates-Oman
borders. The possibility that they might have been sexual
dimorphs is discounted because there are no features in
common between the two. For instance, the change in shape
on the final whorl of the smaller species does not occur on the
larger species.

The nautiloids contrast with those described by Noetling
from the Late Cretaceous of the Mari Hills (Pakistan), where
species of Cimomia and Deltoidonautilus do not seem to
occur (Noetling 1897).

There seems to be little change in the relatively low
nautiloid diversity after the end of the Cretaceous. The
apparent loss of Cymatoceras is matched by the increase in
the Hercoglossidae, mostly in the form of the new suture-line
pattern of Hercoglossa. This change in shell architecture
could represent a wider range of depth tolerance within the
superfamily.

SYSTEMATIC DESCRIPTIONS

Superfamily NAUTILOIDEA de Blainville, 1824
[Name elevated from family rank, Shimanskiy, 1957]
Family HERCOGLOSSIDAE Spath, 1927

The distinction between some of the genera in this family is
rather dubious: Cimomia and Deltoidonautilus in particular
seem to overlap. Kummel (1964, K456) suggests that Cimo-
mia is ‘a morphologically transitional form between Eutre-
Phoceras and Hercoglossa’. Both Cimomia and

| Deltoidonautilus, if indeed they are distinct, seem to fore-

shadow Nautilus itself in their shell morphology.

Genus DELTOIDONAUTILUS Spath, 1927

TYPE SPECIES. Nautilus sowerbyi J. de C. Sowerby, 1843 (see
footnote by Kummel, 1964: K456-57), from the Lower
Eocene, London Clay.

SYNONYMY. Deltoidonautilus may yet prove to be a junior
synonym of Angulithes de Montfort, 1808, because of the
various intrepretations of the type species of Angulithes,
Nautilus triangularis de Montfort, 1802. Kummel accepted
the name in 1956 but rejected it as a nomen dubium in the
Treatise (1964). It is not clear in de Montfort’s (1808: 8)
description, whether the type specimen came from the Lower
Kimmeridgian or from the Cenomanian. It is beyond the
scope of the present work to establish the type material of this
taxon, with the result that Kummel’s (1964: K456) later
opinion that Angulithes should be treated as a nomen dubium
is followed here.

DESCRIPTION. Characteristically smooth, involute and often
compressed with a sub-carinate venter in early to middle
growth stages. Suture sinuous and projected forwards on the
venter. Distinguished from Cimomia by the carinate venter.

RANGE. Upper Cretaceous (Cenomanian) to Oligocene; cos-
mopolitan.

Deltoidonauitilus salisfilius sp. nov. Plate 1, figs 1-3

21861 Nautilus rota Blandford: pl. 25, fig. 2 only.
1928 Nautilus jordani Zittel; Lees: 663, pl. 44, fig. 6.

HOLOTYPE. BM C.59590, from the ‘Main Gastropod Bed’,
bed 6, at Jebel Bu Milh (JBM 2), NNW of Al Ain; base of
Simsima Formation, Maastrichtian, probably Lower Maas-
trichtian.

OTHER MATERIAL. Thirteen paratypes from the same locality
and horizon, BM C.59591—603, Morris and Smith Collection.
Three additional paratypes apparently from the same locality
and horizon, G.M.Lees collection (mentioned Lees, 1928),
BM C.31034-36. Two specimens from the basal 2 meters of
the Simsima Formation at Jebel Faiyah (JF2), U.A.E., BM
C.59607-08. ?Lower Maastrichtian, Morris and Smith Collec-
tion. One specimen from Jebel Faiyah (JF1b), loose, but
probably from the basal Simsima Formation, ?Lower Maas-
trichtian, Gale, Morris and Smith collection, BM C.59609.
One specimen from Jebel Buhays (JB1b) also loose but
?basal Simsima Formation, Gale, Morris and Smith collec-
tion, BM C.59610. All the specimens found apparently occur

252

just above the flooding surface at the base of the Simsima
Formation, probably at a similar horizon to Libycoceras sp.
and Nostoceras spp.

DESCRIPTION. Shell rather smail, the two near complete
specimens from Jebel bu Milh are both approximately 102
mm in diameter. Shell planospiral, smooth, involute, non-
umbilicate on outer shell surface; internal mould with a
narrow, shallow umbilicus. Compressed, discoidal, sub-
oxycone with rounded to subcarinate venter on inner and
middle whorls; outer whorl, ie. body-chamber, rounded and
much less compressed, occupying approximately one third of
the last whorl. Siphuncle well dorsal of centre. Suture with
well-rounded but relatively narrow ventral saddle that bends
forward towards the aperture in a way that is more prominent
in the sub-oxycone stages than in the more rounded whorls
where the few final sutures occur; broad, shallow, sub-
symmetrical lateral lobe; relatively low, smallish lateral
saddle close to the umbilical margin. Sutures show consider-
able crowding towards the body-chamber in what seems to be
a fully grown individual. Siphonal sinus moderate on the
penultimate whorl, observed only on specimen BM C.31034.

COMPARISON WITH OTHER SPECIES. Nautilus fleuriausianus
d’Orbigny (1840: 82, pl. 15) has a rounded venter similar to
the body-chamber of the present species, but does not have
the earlier, more oxycone stage. It is apparently of Upper
Cenomanian age from the ‘craie a Caprines’ at Ile Madame,
Charente, France. It may prove to be a more rounded
specimen of the widespread species named Nautilus triangu-
laris (de Montfort; d’Orbigny, 1840: pl. 12; ?=Angulithes
triangularis de Montfort 1808). Two nearly complete speci-
mens from the Cenomanian of Sidmouth (BM C.931), and
from the Upper Grey Chalk near Folkestone (BM C.8320),
are both carinate and show no sign of change to a round
ventered body-chamber at approximately 160 mm and 180
mm diameter respectively. Nautilus mermeti Coquand, 1862,
from Algeria, may also prove to be a synonym of this
Cenomanian species.

Nautilus westphalicus Schliiter, 1876 from the Late Santo-
nian or Lower Campanian Quadraten Kreide from Dulmen
in Germany also has a carinate body-chamber. Nautilus galea
Fritsch, 1872, from the Iserschichten in Bohemia, has a
carinate venter on the body chamber, has much broader
whorls and apparently has less sinuous sutures.

There may be three separate taxa amongst Blanford’s
(1861) illustrations of his species Nautilus rota: his pl. 24 fig. 3
and pl. 25, fig. 1 clearly belong to Cymatoceras, while the
smooth internal mould of his pl. 25, fig.2 has the flexed suture
typical of Deltoidonautilus and could well belong in the
present species.

Genus CIMOMIA Conrad, 1866

TYPE SPECIES. Nautilus burtini Galeotti, 1837, from the
Eocene of Belgium (a very similar species to C. imperialis (J.
Sowerby, 1812)).

N.J. MORRIS

Cimomia aff. sowerbyana (d’Orbigny, 1840) Plate 1,
fig. 4; Plate 2

Nautilus sowerbyanus d’Orbigny: 83, pl. 16.
Nautilus sowerbyanus d’Orbigny; d’Orbigny: 189.
Nautilus jordani Wanner: 143, pl. 19, fig. 21.
Cimomia jordani (Wanner); Kummel: 451.

aff. 1840
aff. 1850
1902
1956

MATERIAL. A single fairly well-preserved specimen, BM
C.59611, from the base of the Simsima Formation at Jebel
Fayah (JF 2), associated with Deltoidonautilus salisfilius sp.
nov.; the body chamber is missing. A single poorly preserved
but virtually complete specimen from Jebel Buhays (JB 1b),
also apparently from the base of the Simsima Formation, BM
C.59612. Three doubtful specimens from Jebel Rawdah (JH
1), one of them in situ in the base of bed 5, BM C.59613, and
the other two loose on the scree below but apparently from
the same horizon, BM C.59614-15.

In his original description, d’Orbigny stated that the local-
ity of the type specimen of Nautilus sowerbyanus was
unknown, but later, in the Prodrome (d’Orbigny, 1850: 189),
he gave the locality as Montrichard, France. The specimen
was in the Musée d’Histoire Naturelle, Paris, the locality
details having been sent by M. Cordier, and it seems quite
likely that Fischer, who organised d’Orbigny’s collection,
must have sorted out the locality before publication of the
Prodrome.

Wanner’s holotype of Nautilus jordani came from the
uppermost white chalk near Bab-el-Jasmund to the north of
Dachel, western Egypt.

DESCRIPTION. Shell medium-sized, planospiral, smooth,
involute. External shell non-umbilicate, with the umbilicus of
the internal mould plugged by a slight recurved extension of
the lateral saddle, almost in the form of a small very shallow
lobe. Shell inflated to spheroidal with flanks converging to
the rounded but slightly flattened venter. Suture with a very
low, gently rounded ventral saddle, a symmetrical, relatively
shallow lateral lobe and a relatively prominent well-rounded
lateral saddle.

COMPARISON WITH OTHER SPECIES. ?Cimomia applanatus
(Wanner, 1902: 143), another species named from western
Egypt, could be another variant of this species but is consid-
erably more evolute than any of the specimens considered
here from the same area. The identification of Nautilus
desertorum Quaas (1902: 299) is also uncertain, because it is
not possible to judge from the original illustration whether
the suture-line is sinuous like that of Cimomia or straighter
like that of Eutrephoceras.

Cimomia aff. sowerbyana is similar in general shape to
Nautilus sublaevigatus d’Orbigny (1850: 189; nom. nov. for
N. laevigatus d’Orbigny, 1840: pl. 17, non Reinecke, 1818).
The first locality listed where d’Orbigny collected this species
is Martrou near Rochefort. He also listed a number of other
localities including Montrichard, Uchaux and several from
Cotentin; these span a range from Turonian to Maastrichtian.
The sutures of N. sublaevigatus seem much less sinuous than
those of Cimomia sowerbyana and suggest that this species

PLATE 1

Figs 1-3 Deltoidonautilus salisfilius sp. nov. Base of Simsima Formation Maastrichtian, ? Lower Maastrichtian, Jebel Bu Mil;ch, NNW of Al
Ain. 1, holotype, BM C.59590; 2, paratype, BM C.59591; 3, paratype, BM C.59592, arrow shows poistion of the siphuncle; all x 1.
Fig. 4 Cimomia aff. sowerbyana (d’Orbigny, 1840). Base of Simsima Formation, ?Lower Maastrichtian, Jebel Fayah, south of Dayid,

ventral view; BM C.59611; x 1.

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’

MAASTRICHTIAN NAUTILOIDS 253

J. MORRIS

254

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MAASTRICHTIAN NAUTILOIDS

belongs to Eutrephoceras. Similarly N. vastus Kner, 1850,
Cimomia cf. forbesi dArchiac & Haime, 1854 (a Danian
species), and Eutrephoceras sphaericus (Forbes, 1846) (holo-
type from Pondicherry, India, BM C.73524; not pre-occupied
by Nautilites sphaericus Martin, 1809), may belong to Eutre-
phoceras.

A number of Late Cretaceous specimens in the BM(NH)
collection from the United States also indicate that Nautilus
dekayi Morton, 1833, does not have the sinuous suture-line of
Cimomia and properly belongs in Eutrephoceras.

REFERENCES

Blainville, H. M. D. de 1824. Dictionaire des science naturelles; Mollusques, 32:
392 pp. Strasbourg and Paris.

Blandford, H. F. 1861. The Fossil Cephalopods of the Cretaceous rocks of
Southern India, (Belemnitidae - Nautilidae). Memoirs of the Geological
Survey of India, Palaeontologia Indica, 1: 40pp., 25pls.

Conrad, T. A. 1866. Observations on Recent and fossil shells, with proposed
new genera and species. American Journal of Conchology, 2: 101-03.

Coquand, M. H. 1862. Géologie et Paléontologie de la région sud de la Province
de Constantine. Marseille.

Forbes, E. 1846. Report on the fossil Invertebrata from southern India
collected by Mr. Kaye and Mr. Cunliffe. Transactions of the Geological
Society of London, (2) 7: 97-174, pls. 7-19.

Fritsch, A. & Schloenbach, U. 1872. Cephalopoden der béhmischen Kreidefor-
mation. 52 pp., 16 pls. Prague.

Galeotti, M. H. 1837. Mémoir sur la constitution géognostique de la province
de Brabant, en réponse a la question suivante: décrire la constitution
géologique de la province de Brabant, déterminer avec soin les éspeces
minérales et les fossiles que les divers terrains renferment et indiquer la
synonymie des auteurs qui en ont déja traité. Mémoires couronneés de
l'Académie Royale des Science et Belles Lettres, 12: 1-192, pls. 1-4. Brux-
elles.

255

Kner, R. 1850. Versteinerungen des Kreidemergels von Lemberg und seiner
Umgebung. Naturwissenschaftliche Abhandlungen (Haidinger), 3 (2): 1-42,
pls 1-S.

Kummel, B. 1956. Post-Triassic Nautiloid Genera. Bulletin of the Museum of
Comparative Anatomy at Harvard College, 114 (7): 321-494, 28 pls.

1964. Nautiloida-Nautilida. Jn Moore, R.C. (editor), Treatise on Inverte-
Brate Paleontology, K, Mollusca 3, 519 pp. The Geological Society of
America and the University of Kansas Press.

Lees, G. M. 1928. Geology and tectonics of Oman and parts of southeastern
Arabia. Quarterly Journal of the Geological Society of London, 84: 585-670.

Montfort, D. de 1808. Conchyliologie systématique et classification méthodique
des coquilles; offrant leur figures, leur arrangement générique, leurs descrip-
tions caractéristiques, leurs noms; ainsi que leur synonymie en plusieurs
langues. 1: 409 pp., 100 pls. Paris.

Morton, S. G. 1833. Supplement to the ‘Synopsis of the Organic Remains of the
Ferruginous Sand Formation of the United States’, contained in Vols XVII
and XVIII of this Journal. American Journal of Science, 23 (2): 288-94.

Noetling, F. 1897. Fauna of Baluchistan, Vol. 3, part 1, Fauna of the Upper
Cretaceous (Maastrichtian) Beds of the Mari Hill. Memoirs of the Geological
Survey of India, Palaeontologia Indica, (16) 1 (3): 79 pp., 23 pls.

Orbigny, A. d’ 1840. Paleontologie Francaise. Terraines Cretacées. Céphalopo-
des. 1: 662 pp., 148 pls.

1850. Prodrome de Paléontologie stratigraphique universelle des animaux
mollusques et rayonnes faisent suite au cours elementaire de Paléontologie. 3
vols. Masson, Paris.

Quaas, A. 1902. Beitrag zur Kenntniss der Fauna der obersten weissen
Kreidebildungen in der libyschen Wuste (Overwegischichten und Blatter-
thone). Palaeontographica, 30 (2) (4): 153-355, pls 20-33.

Schliiter, C. 1876. Cephalopoden der oberen Kreide. Palaeontographica, 24
(2): 123-263, pls. 36-55.

Shimanskiy, V. N. 1957. Sistematica i Filogenia otryada Nautilida. Bulletin de
la Societe des Naturalistes de Moscou, Geologie, 32 (4): 105-20 [In Russian].

Sowerby, J. & Sowerby, J. de C. 1812-1846. A Mineral Conchology of Great
Britain. Vols 1-7, 650 pls.

Wanner, J., 1902. Die Fauna der obersten weissen Kreide der libyschen Wiste.
Palaeontographica, 30 (2) (3): 91-151, pls 13-19.

Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):257-265

Issued 30 November 1995

Maastrichtian Inoceramidae from the United
Arab Emirates-Oman border region

NOEL J. MORRIS

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD

Synopsis. Descriptions of Inoceramids from Upper Cretaceous post-Semail Nappe emplacement deposits include
Endocostea (Selenoceramus) semaili sp. nov. Most specimens are from the Simsima Formation in Jebel Rawdah,
immediately below the massive limestone facies, and they indicate a mid-Maastrichtian age.

INTRODUCTION

Small collections of Inoceramidae from two localities on the
western edge of the Omani Mountains, close to the border
between the United Arab Emirates and Oman, are preserved
largely as uncrushed internal moulds. The first is from the
Simsima Formation in Jebel Rawdah just north of the Al Ain
to Hatta Road, about 10 Km east of the Madam Roundabout,
just east of the Omani Border Post, and is of mid-
Maastrichtian age. The second is from the Loftusia-rich marls
below the Simsima Formation in Jebel HuwaYyah (known as
Fossil Valley), to the east and north-east of Al Ain, and is
apparently of Lower Maastrichtian age. It is quite clear that
the specimens from the higher horizon, i.e. in Jebel Rawdah,
assist with the age determination of the strata, and they may
have potential for international correlation.

The Inoceramidae show that at least three distinct lineages
existed in the eastern Arabian area in mid-Maastrichtian
times. Each lineage seems to fall in some part of Whitfield’s
(1877) genus, Endocostea, which is in much need of revision.
Any serious revision is, however, beyond the scope of this
paper. Subgeneric names are used tentatively until we have a
better understanding of the phylogeny of the group. The
apparent lack of specimens of Trochoceramus, which is
widespread across Africa in late Campanian and Maastrich-
tian times, may reflect the palaeoecology. The water may
have been either too shallow or too warm, or the sediment
too coarse.

The specimens described here were collected by Drs Gale,
Morris and Smith in January 1992, by Drs Nolan and Skelton
during an earlier visit (Nolan et al. 1990), and by members of
the Emirates Natural History Group.

Class BIVALVIA Linnaeus 1758
Subclass PPERIOMORPHIA Beurlen, 1944
[nom. trans. Newell, 1965]
Superfamily INOCERAMOIDEA Giebel, 1852
[?=Ambonychioidea Miller, 1877]

| : ‘ ‘ si
The Inoceramidae share with Ambonychia the thick calcitic
_ inter-umbonal ligament area not present in adult Pterioidea.

Family INOCERAMIDAE Giebel, 1852

DESCRIPTION. Variously shaped, concentrically lamellose or
plicated or radially plicated Pteria-like bivalves which some-
times grew to a large size. The outer calcareous shell layer
consists of vertical calcite prisms which are modified to
pyriform laths on the ligament area. The ligament, usually
multivincular, is attached largely to this external layer; inner
shell layer nacreous; usually lacking hinge teeth; usually
equivalve or subequivalve, some species inaequivalved. A
complex of muscle scars close to the umbones is consistant
with byssal fixation. Apparently monomyarian with the pos-
terior adductor often close to the posterior ventral margins
but often obscure. It is very difficult to be certain whether an
isolated muscle scar situated about halfway below the
umbones and towards the anterior margin is a large pedal
muscle scar or a small anterior adductor. The Inoceramidae
became extinct by the beginning of the Tertiary.

COMMENTS. The ligament area of these later Cretaceous
species is seldom well-exposed and the pits characteristic of
earlier species have only been observed on one of the species.
It remains possible that lineages other than Tenuipteria
(Dhondt, 1983b) have no pits for the differing ligament types.

THE INTERNAL SHELL RIB. Many post-Coniacian Inocerami-
dae have an internal shell rib, sillon or Hohlkehle. The nature
of this internal rib has been discussed on a number of
occasions (especially Seitz, 1967: 14-41). It occurs commonly
in a number of Inoceramidae from at least as early as the
Santonian. It is here considered to be an architectural charac-
ter of the shell, and it is clearly variable in some of the taxa
that have it. Two specimens of Endocostea sp. from the
Santonian-Campanian, Haslam Formation of Brannan
Creek, Nanaimo District, British Columbia, Canada, demon-
strate this point; in BM LL 28194 the inner ‘rib’ is poorly
developed, starting at approximately 6 cm from the umbo of
the right valve and continuing for only 2 cm, whereas BM LL
28217, a slightly smaller individual from the same locality and
horizon, developed the rib at 1-5 cm from the umbo of the
right valve, from which position it continues for at least 4-5
cm towards the posterior ventral margin.

In three well-preserved specimens from the late Campa-
nian, Fort Pierre Group (two examples of ‘/noceramus? aff.
barabini Morton, BM L 21569 from the Cheyenne River
Section, BM L 7577 from the Powder River Range, Montana,
and a single example of ‘/noceramus’ tenuirostris Meek &

258

Hayden, 1862, BM L 21571, from the Cheyenne River), no
fully developed internal rib is present. In its place are raised
ridges on the innermost shell following the track of the
normal position of the inner rib. These end distally as
tangents to the 1 cm wide muscle scar that has been referred
to as the posterior adductor. Clearly, inoceramids have an
unusual arrangement of their soft parts, but it would appear
the inner rib is formed as a sensitive emplacement of shell
material to enhance strength in the part of the shell under
stress from muscular shell closing.

The internal rib is also present on some Jurassic species; a
well-preserved specimen of Parainoceramus ventricosus (J. de
C. Sowerby, 1823) from the Lower Jurassic, Pliensbachian, at
Brockthorpe, near Gloucester, shows precisely the same
details as ‘Inoceramus’ barabini, with an internal rib running
down to the adductor. Some Campanian specimens of
Endocostea, however, seem to show the internal rib running
very close to the ventral margin, apparently past the position
of the adductor.

There are also preservational problems in observing the
internal rib. Early diagenesis of the aragonite shell interior
can destroy the rib, and there are many examples of inocera-
mids preserved only as internal moulds of the outer calcitic
shell layer.

The rib is nacreous, is often hollow and has an arch-shaped
section. In all the examples that I have examined where two
valves are preserved together, one rib is an almost perfect
mirror image of the other. The rib is in a morphologically
comparable position in all species in which it occurs. It shows
distinct evolutionary changes, one of the most notable being
extreme broadness in some species of Seleniceramus (Seitz
1967), though it is usually distinctly narrow. The internal rib
may be close to a muscle scar at its distal end.

The claim that the internal rib is formed by, or in response
to, a parasite may explain why it is sometimes filled with
matrix. This can, however, be equally well explained if it is
due to the activity of shell borers: if a hole is bored from the
outside of the shell into the hollow of the rib, the damage
cannot be repaired by deposition of more shell material as it
is separated from the outer surface of the mantle by the
thickness of the rib itself. The same would be the case if the
inoceramid was already dead when its shell was drilled. All
the cases of matrix- filled inner ribs that I have examined
show evidence of drilling from the shell exterior.

The inner rib is clearly of some importance for taxonomy at
both the species and higher levels. Its variability in some
species and its possible diagenetic loss mean that it needs to
be used with caution.

Genus ENDOCOSTEA Whitfield, 1877

TYPE SPECIES. E. typica Whitfield, 1877, by original designa-
tion. The lectotype (USNM 12261) was selected by Seitz
(1967: 54-55, pl. 2, fig. 4). It is the example ‘c’ of Whitfield
(1880: pl. 9 fig. 3) and is from the Pierre Shale at Old Woman
Fork on the Cheyenne River, Black Hills, South Dakota,
which is of late Campanian age according to Cobban &
Reeside (1952: 1011) (non Inoceramus cripsi var. typica
Zittel, 1866: 98, from the late Cretaceous of Grunbach in
Neuen Welt and the Gosau Valley, Austria).

DisCUSSION. The generic name was introduced for relatively
small inoceramids with a well-developed inner rib radiating

N.J. MORRIS

from the umbo towards the posterior ventral margin but
stopping well short of that margin. The species is otherwise
very similar to Whitfield’s own interpretation of Inoceramus
barabini Morton (Whitfield, 1885: 75-76), except that the
latter species apparently does not have the internal rib.

Subgenus ENDOCOSTEA Whitfield, 1877

A number of species or subspecies of Endocostea are rather
convex with prominent or enrolled anterior umbones. These
occur in the late Campanian and Maastrichtian and resemble
Endocostea (Endocostea) coxi (Reyment). They include Jnoc-
eramus balticus pteroides (Giers, 1964: pl. 1, fig. 6) from the
Upper Campanian, Polyplocum Zone of Haldem, Westfalia,
and according to Sornay (1976) from Dau, Charente. They
possibly also include Inoceramus borilensis Jolkicev, said to
be from the Maastrichtian in Bulgaria. However, specimens
that are very similar to J. bakalovi (said by Jolkicev (1961) to
occur at the same horizon) occur with ammonites of the
Upper Campanian, Donetzianum Horizon, in Nigeria. They
also include Inoceramus impressus d’Orbigny (lectotype,
Museum d@ Histoire Naturelle, Paris, 7592a, figured Sornay,
1957: 129) from the ?Upper Campanian of Royan, and
‘Inoceramus impressus’ d’Orbigny (pars, but not including
the lectotype) from the Maastrichtian of Cotentin, Manche,
France. These are provisionally included in the subgenus
Endocostea sensu stricto, although their relationship with the
less convex type species remains to be confirmed.

Endocostea (Endocostea) coxi (Reyment, 1958) Plate 1,
figs 24

1958 Inoceramus coxi Reyment: 140, pl. 3, figs 4-6.

MATERIAL. The type material comes from Auchi in north-
central Nigeria and is preserved in a medium grained, slightly
ferrugenous sandstone (holotype BM L 82963). Some better
preserved specimens occur on the [kot Ekpene Road in the
Calabar area associated with magnificent specimens of Tro-
choceramus ianjonaensis and are therefore of ‘mid’ (probably
low Upper) Maastrichtian age. Another specimen in the
BM(NH) collection comes from Madagascar. Three speci-
mens were collected by Nolan and Skelton from the south-
west face of Jebel Rawdah, BM LL 4164749.

DESCRIPTION. Convex, equivalved species with strong radiat-
ing sulcus posterior to the umbones, delimiting a much less
convex posterior area. Somewhat quadrate, but with
umbones well to the anterior and prosocline. Strong radiating
internal rib well to the posterior of the convex part of the
shell. Strong but low rounded comarginal ribs, separated by
wider interspaces on anterior and flank, ie. the convex part of

the shell, fading posteriorly. Smoother in later growth stages. |

COMMENTS. This species was included in Cordiceramus by
Dhondt (1983a) following Seitz (1967), a genus that was
included as a synonym of Haenleinia by Cox (1969). Cordice-
ramus Seitz (1961: 110, ex. Heinz, 1932) has a very distinctive

wide external radiating furrow from the posterior of the —

umbo to the posterior ventral margin and is more or less
equivalve. The type species, C. cordiformis Sowerby, is a
relatively tall species with a short hinge line, whereas Haen-
leinia is an elongate genus related to Endocostea, with a
posterior shell twist, resulting in an inequivalve shell in that

MAASTRICHTIAN INOCERAMIDAE

PLATE 1 Fig. 1 Endocostea (Endocostea) sp. indet.; BM LL 41651, left valve; south-west face of Jebel Rawdah, near base of Simsima
Formation, ‘mid’-Maastrichtian; Skelton & Nolan Collection, x 2.

Figs 2-4 Endocostea (Endocostea) coxi (Reyment). 2, BM LL 41647, 3, BM LL 41648, both internal moulds of a right valves; 4, BM LL
41649; internal mould of a left valve; all from the south-west face of Jebel Rawdah, Nolan and Skelton Collection, x 2.

Figs 5,6 Endocostea (Selenoceramus) semaili sp. nov.; 5, BM LL 41639, holotype, Jebel Rawdah, section 1, from scree below bed 5,
‘mid’-Maastrichtian, Gale, Morris & Smith Collection, x 1; 6, BM LL 41640, paratype, Jebel Rawdah, section 1, from bed 5,
‘mid’-Maastrichtian, Gale, Morris & Smith Collection, x 2.

259

260

area. The radiating furrow in some specimens of E. coxi is
found on composite moulds and reflects its internal rib.
Well-preserved specimens of E. coxi from near Calabar,
Nigeria, have no external furrow and do not seem properly
placed in either Cordiceramus or Haenleinia.

Very similar specimens occur in the low Upper Maastrich-
tian part of the St Lucia Formation in South Africa (W.J.
Kennedy, personal communication).

Plate 1, fig. 1

1844 Inoceramus impressus d’Orbigny (pars): 515-16, pl.
409.

Inoceramus (Endocostea) impressus
Tsankov et al.: 97, pl. 42, fig. 2.

Endocostea (Endocostea) sp. indet.

21981 d’Orbigny;

MATERIAL. A single specimen from the south-western face
of Jebel Rawdah, low in the Simsima formation, Skelton and
Nolan Collection, BM LL 41651, ‘mid’-Maastrichtian.

THE TYPE MATERIAL OF INOCERAMUS IMPRESSUS. Sornay
(1957) chose the specimen from Royan in the d’Orbigny
Collection as lectotype. This seems to have a much more
prominent umbo than the present species, more closely
resembling E. borilensis Jolkicev and E. coxi Reyment. The
specimen here (Plate 1, fig 1) although not well preserved, is
much more like the species from Cotentin that resembles the
left valve only of d’Orbigny’s figure. Specimens from the
Maastrichtian of Cotentin in the de Gerville Collection in The
Natural History Museum, resembling the right valve in
d’Orbigny’s figure, seem more closely related to E. coxi and
are mentioned under Endocostea (Endocostea) above.

DESCRIPTION. Small elongate species with internal rib.
Umbones well to the anterior but not prominent and incurled
like the contempory species E. coxi. By means of this
character, specimens from the Upper Maastrichtian of Cote-
ntin are easily separable from £E. coxi, although in
d’Orbigny’s description the species was described as inaequi-
valve. It seems that he considered that specimens with very
prominent umbones, here thought to be related to E. coxi,
were right valves of the species described here. The species
also seems to occur commonly in the Maastrichtian rocks of
the St. Lucia Formation from Zululand (W.J.Kennedy Col-
lection).

Subgenus SELENOCERAMUS Seitz, 1967 (ex. Heinz, 1932)

TYPE SPECIES. Inoceramus (Selenoceramus) selenae Seitz,
1967, p. 95, by original designation.

Heinz’s definition of Selenoceramus does not contain char-
acters that distinguish this genus from others, except in his
description of the two new species included in the genus. His
type species, Selenoceramus pulcher, is treated by Seitz (1967:
94), somewhat dubiously, as a nomen nudum, apparently
because there was no stated type specimen. Seitz attempted
to validate the genus in terms of Heinz’s original intention.
Because of the ambiguity in ICZN article 13(a), it may prove
necessary to apply to the International Commission to deter-
mine whether the genus should date from Heinz (1932) or
Seitz (1967).

COMMENTS. Selenoceramus includes some rather small
rounded inoceramids with concentric sculpture that show a
marked change in growth direction following an evenly

N.J. MORRIS

curved young stage and have a wide internal rib. The material
from the Omani Mountain area shows that this subgenus
continues well into the Maastrichtian.

Plate 1,
figs 5, 6

21865 Inoceramus cripsi Mant, var. regularis d’Orbigny;
Zittel: 93-99, pl. 14, fig. 3 only.

21974 Inoceramus regularis d’Orbigny; Blank et al.: 85, pl.
Pl, Wis Ze joll, BI, ties, Al.

HOLOTYPE. BM LL 41639, from scree below bed 5, Simsima
Formation, Maastrichtian; Jebel Rawdah, section 1; Gale,
Morris and Smith Collection.

Endocostea (Selenoceramus) semaili sp. nov.

OTHER MATERIAL. Two small paratypes, BM LL 41640-41,
from the same horizon and locality as the holotype; Gale
Morris and Smith Collection. Three further paratypes, BM
LL 4164244, from the south-western outcrop in Jebel Raw-
dah, apparently from the same horizon; Nolan and Skelton
Collection.

AGE. The accompanying ammonites and other species of
Inoceramidae suggest that this species is of low Upper
Maastrichtian age.

DESCRIPTION. Medium-sized inoceramids with close, regu-
lar, rounded, concentric ribs (which show a tendency to
straighten) parallel to the anterior-ventral margin. Ribs
approach dorsal margin at a relatively high angle to the
posterior of the umbones. Maximum shell height below
posterior point of hinge. Umbones orthocline, towards but
not at anterior. Shells gently convex to a shell height of 70
mm in holotype, then a sharp change of direction to give
increased width. This change of shell curvature is at variable
distance from the umbones and occurs at an earlier growth
stage in two of the paratypes.

The inner rib is angled at about 45° to the hinge line, and is
only represented by a trace in the holotype. Two of the
paratypes (LL 41642, 43) have a poorly preserved but distinc-
tively wide and shallow internal rib, typical of the subgenus
Selenoceramus.

The convexity and style of ribbing of E. semaili matches
that of E. cf. semaili (Fig. 1) from the Lower Maastrichtian,
Acanthoscaphites tridens Zone, at Nagoryani, Ukraine,
except that it is smaller and the ribbing is consequently closer.
However, approximately the same number of ribs are
present. Unfortunately, specimens from Nagoryani are not
well-preserved on the inner shell surface, making it very
difficult to interpret the form of their interior ribs.

The species should be compared with Endocostea mandem-
bataensis (Sornay, 1973: 90-91, pl. 4, fig. 4; Fig. 4) from
Mandembata, southern Madagascar, ‘Lower and Middle’
Maastrichtian. Unfortunately, this species is defined on a
single rather incomplete figured specimen. The anterior
portion of the shell, a left valve, is missing, which does not
allow identification at the species level. This species is a
nomen dubium until further material from the type locality 1s
described. Sornay himself (1973: 11) stated that the material
was not sufficiently well preserved to enable him to work out
its relationships.

Endocostea semaili should also be compared with
Endocostea kneri Boehm (1909: 53; nom. nov. for Inocera-
mus impressus Kner, 1850, non d’Orbigny) from Nagoryani,
Ukraine, presumably from the Lower Maastrichtian “Tridens

MAASTRICHTIAN INOCERAMIDAE

Fig. 1 Endocostea (Selenoceramus) cf. semaili sp. nov; ‘Tridens’
Zone, Lower Maastrichtian, Nagoryani, Ukraine; right valve; M.
Mackay Collection, BM LL 41653, x 0-5.

Fig. 2 Endocostea ?(Cataceramus) sp. indet.; “Tridens’ Zone,
Lower Maastrichtian, Nagoryani; M. Mackay Collection, BM LL
41654, x 0-5.

_ Zone’. When compared with sixteen specimens from Nagory-
ani in the collections of The Natural History Museum, it is
fairly certain that the type of this nominal species is badly
distorted. It has become much more elongated than all of
them except one. The latter specimen, BM LL 61647, has a
normal, rather rounded, right valve, but diagenetic distortion
has decreased the height of the left valve. The similarity of
the form of the sculpture of this left valve with that of Kner’s
figure suggests that his specimen was also similarly distorted.
For this reason I consider that Endocostea kneri is a nomen
dubium until such a time as its true shape can be described,
possibly when the Inoceramidae from Nagoryani are revised.
Specimens from Nagoryani are illustrated in Figs 1 and 2.
Endocostea semaili sp. nov. is very similar to E. (Catacera-
| mus) baltica (Boehm, 1909; see Geirs, 1964: pl. 1, fig. 1 only)

261

from the late Santonian (? and Lower Campanian) of Ger-
many, but it differs in having the umbones slightly more
posterior. Also the comarginal ribs are much more regularly
curved in Boehm’s species, giving it a very rounded appear-
ance, whereas there is a slight straightening of the ribs ventral
to the umbones in E. semaili. The nature of the internal rib of
E. (Cataceramus) baltica is in doubt.

Inoceramus sornayi Dhondt (1993) was a nom. nov. for
Inoceramus regularis d’Orbigny, 1846, non Munster, 1840;
the lectotype was designated by Sornay (1962), and other
specimens were figured by him (Sornay, 1976: pl. 2, figs 3, 4;
pl. 3, fig. 4). 7. sornayi is from the late Campanian and has a
wide internal rib, and like Endocosta semaili, is better placed
in Selenoceramus.

E. (Selenoceramus) semaili has its umbones further from
the anterior margin than E. (S.) sornayi and has a differently .
shaped anterior; its anterior ventral margin has a diagonally
cut away appearance in contrast to the rounded anterior of E.
sornayi (see Sornay, 1962: pl.7, fig. 3, the lectotype; and
Dhondt, 1993: pl.6, fig. 3). E. (S.) semaili may have been a
descendant and chronological subspecies of E. (S.) sornayi. It
has not been found amongst the many Inoceramids in the St.
Lucia Formation in South Africa, possibly because it is
restricted to more northerly areas.

Subgenus CATACERAMUS Cox, 1969 (ex. Heinz, 1932
?nom. nud.)

TYPE SPECIES. Inoceramus goldfussianus d’Orbigny, 1845, by
original designation.

Cox’s designation of Inoceramus goldfussianus d’Orbigny
as the type species is unfortunate because it is not the same as
I. balticus Boehm as he claimed. More unfortunately, the
specimen he figured as /. balticus has been separated from the
type of that species by Giers, and is in fact the holotype of /.
marki (Giers).

Heinz (1932: 15) proposed Cataceramus, with type species
Inoceramus balticus Boehm, as a subgenus of Selenoceramus.
Cox implied that Heinz’s description does not fulfill the
requirements of Article 13 (a) (i) in that there is no defini-
tion, and the name may therefore not be valid from that date.
The type species, /. balticus, was emended by Giers (1964:
238-39). As pointed out by Dhondt (1993), Cox (1969)
wrongly included /. balticus as a synonym of I. goldfussianus,
and therefore changed the nature of the genus. Athough
Heinz (1933) went on to use his generic name he still did not
include characters that distinguished Cataceramus from other
genera. Seitz (1967: 49) dismissed Cataceramus Heinz, 1932,
as a synonym of Endocostea Whitfield.

IDENTITY OF THE TYPE SPECIES. Inoceramus goldfussianus
d’Orbigny, 1845, was unnecessarily emended by Sornay
(1957; 1976) to I. goldfussi. The species was described from
the late Cretaceous of Royan by d’Orbigny, and he included
the specimen figured by Goldfuss (1836: 116, pl. 112, fig. 4)
as Inoceramus cripsii Mantell in his synonymy. Sornay (1957)
figured the lectotype (d’Orbigny collection, no. 7593), previ-
ously chosen by Heinz, and also some new specimens (Sor-
nay, 1976: pls 4, 5).

COMMENTS. In spite of the difficulties with the identity of the
type species, this is the ‘Jnoceramus balticus’ group of many
authors. Although Dhondt (1993) suggested that no close
relative of C. goldfussianus can be recognised, both the

262

lectotype and Sornay’s, 1976, pl. 4, fig. 2, have a definite
indication of an internal rib in addition to a general similarity
in shape to J. balticus. The clear difference is the more spaced
out nature of the ribbing, which at present I regard as only
significant at species level. Endocostea (Cataceramus) gold-
fussianus would seem to be of late Campanian age as under-
stood here. Specimens from Celles near Riberac, Dordogne,
France, suggest that the species occurs as early as the Vari
Zone, Upper Campanian.

Upper Santonian and Lower Campanian species of Catac-
eramus are extensively described and illustrated by Seitz
(1967) under the subgenus Endocostea sensu stricto. The
inner rib of Seitz’s E. baltica baltica is apparently intermedi-
ate in form between Selenoceramus and Endocostea sensu
stricto. Unfortunately, the inner shell layer is not preserved
on the lectotype of Bohm’s species (Giers, 1964: pl. 1, fig. 2).

A number of additional species of this subgenus recognised
in the Upper Campanian and Lower Maastrichtian are in
need of further revision, but this is beyond the scope of the
present paper. Two subspecies are, however, of stratigraphi-
cal value: Endocostea baltica baltica Boehm (1907: 113; 1909:
pl. 11, fig. 2; emended Giers, 1964: 238-39, pl. 1, figs 2-4)
and E. baltica marki occur in the Upper Santonian of
Dulmen. Giers (1964), however, pointed out that E. baltica
ranges higher, and he considered the subspecies E. baltica
haldemensis (Giers 1964: 243-44, pl. 2, fig. 2) from the
Polyplocum Zone of Haldem, Lemforde, Westfalia (?non
Inoceramus haldemensis Heinz, ?nomen nudum) to be typical
of the Upper Campanian. Two other subspecies, E. baltica
ellipticus and E. baltica sublaevigatus, occur in the low Upper
Campanian at Tercis, south-west France (Dhondt, 1993).

Endocostea (?Cataceramus) aff. goldfussianus
(d’Orbigny, 1846) Plate 2, fig. 4

1846 Inoceramus goldfussianus d’Orbigny: 517, pl. 411.
1976 Inoceramus goldfussi d Orbigny; Sornay: 9-11, pl. 4;
pl. 5, figs 4, 5.

MATERIAL. A single specimen, kindly donated by Mrs Vale-
rie Chalmers and other members of the UAE Natural History
Group, BM LL 41645. Apparently from Jebel Rawdah, its
calcarenite matrix is rich in Lepidorbitolina sp., and it seems
to come from the same horizon as the other material from this
locality.

COMMENTS. Whereas this individual might be a variant of E.
(S.) semaili sp. nov., it has a rather different ribbing pattern,
clearly similar to that of the earlier FE. golfussianus
(d’Orbigny), but closer packed. There is a slight change in
shell curvature suggesting that it would have been a consider-
ably smaller individual when full grown than the typical late
Campanian specimens of western France. It also closely
resembles a specimen from the Lower Maastrichtian,
‘Tridens Zone’, of Nagoryani, Ukraine (Fig. 2).

PLATE 2

N.J. MORRIS

ENDOCOSTEA BEBAHOAENSIS AND RELATED SPECIES. A num-
ber of inflated species of late Campanian and Maastrichtian
age seem to form a natural group. They have umbones well to
the anterior and show no change of growth direction as they
approach full size. They include:

1. Inoceramus cripsi Mantell var. decipiens Zittel, 1865:
95-99, pl. 15, fig. 1; Gosau Beds, Grunbach, Austria.

2. Inoceramus bebahoaensis Sornay, 1973: 89, 90, pl. 3, figs
1, 2, text-fig. 4.

3. Inoceramus balticus beckumensis Giers, 1964: pl. 2, fig. 1,
from the low Upper Campanian, Beckumer Schichten,
near Beckum, Germany.

4. ?Inoceramus balticus Boehm, Blank et al. 1974: 83, pl. 22,
fig. 2.

5. Inoceramus borilensis dauensis Sornay, 1976: 5, 6, pl. 1,

iy, Se ol, A, whys tl, 2.
. Endocostea flexibaltica (Seitz); Dhondt, 1993: pl. 4, fig. 4.
. 2Inoceramus borilensis Jolkicev, 1962: 145, pl. 7.

There is apparently no subgeneric name for this group, but
I consider it inadvisable to introduce one here because the
full relationships within Endocostea are insufficiently under-
stood.

‘Endocostea’ bebahoaensis (Sornay, 1973) Plate 2, figs
1,2

1973. Inoceramus bebahoaensis Sornay: 89-90, pl. 3, figs
1-2, Fig. 4.

MATERIAL. A single well-preserved specimen from the
south—west face of Jebel Rawdah, Nolan and Skelton Collec-
tion, BM LL 61646.

MATERIAL FROM OTHER LOCALITIES. A single specimen, BM
L 74737, from the Upper Maastrichtian, Calcaire a Baculites,
Cotentin Peninsula, Normandy, in the de Gerville (ex J. de
C. Sowerby) Collection. Other specimens are known from
the Maastrichtian of Ianjona and Bebahoa, southern Mada-
gascar (Sornay, 1973), and the St. Lucia Formation, Zululand
(W.J.Kennedy Collection, Oxford University Museum).

DESCRIPTION. Convex medium-sized species of Inocerami-
dae with unchanging growth curvature and only gently coars-
ening, regular, low, rounded, concentric ribs. Umbones well

]

to the anterior and gently prosocline. Our single right valve | _

from Jebel Rawdah is an internal mould and has a shallow

}

narrow groove running from the umbo towards the posterior | _
ventral margin as far as the shell is preserved. This is the |
impression of the internal rib, which does not seem to have 1

been preserved on other specimens.

Figs 1,2 ‘Endocostea’ bebahoaensis (Sornay). 1, BM L 74737, Calcaire a Baculites, Upper Maastrichtian, Cotentin Peninsula, France, de
Gerville Collection; left valve, la, anterior view, 1b, lateral view, x 1. 2, BM LL 41646, south-west face of Jebel Rawdah,
‘mid’-Maastrichtian, near base of Simsima Formation; Nolan & Skelton Collection; right valve, 2a, lateral view, 2b, anterior view, 2c,

dorsal view; X 1.

Fig 3 ‘Endocostea’ cf. bebahoaensis (Sornay). BM LL 41652, Loftusia-Beds, Jebel Huwayyah, ?Early Maastrichtian, Gale, Morris & Smith

Collection, anterior view of incomplete right valve, x 1.

Fig. 4 Endocostea (?Cataceramus) aff. goldfussianus d’Orbigny. BM LL 41645, possibly from near the base of the Simsima Formation, Jebel
Rawdah. ?Maastrichtian, Mrs. V. Chalmers Collection; 4a, lateral view of right valve, 4b, anterior view, x 1.

263

MAASTRICHTIAN INOCERAMIDAE

264

‘Endocostea’ cf. bebahoaensis (Sornay, 1973). Plate 2,
fig. 3

cf. 1973 Inoceramus bebahoaensis Sornay: 89-90, pl. 3, figs

1-2, Fig. 4.

MATERIAL AND COMMENTS. A single fragment, BM 41652,
from the Loftusia-Beds at Jebel Huwayyah, consists of only
the anterior portion of the shell. The anterior position of the
umbo, the steep drop from the convex flank to the anterior
margin and the style of ribbing, demonstrate that this frag-
ment is very similar to Endocostea bebahoaensis, particularly
the specimen from Cotentin (Plate 2, fig. 1j. Ammonite
evidence suggests that the Campanian-Maastrictian boundary
may lie within the Loftusia-Beds, and the occurrence of
Pachydiscus dossantoi (Maury, 1930) in the upper part of
these beds suggests that the age here is at least as high as
mid-Maastrichtian.

Genus PLATYCERAMUS Seitz, 1961, p. 54
(ex. Platyceramus Heinz, 1932, p. 10, nomen nudum)

TYPE SPECIES. Inoceramus mantelli de Mercy, 1877, by origi-
nal designation.

As originally introduced by Heinz (1932: 10), with the type
species designated as ‘Genotyp: Jnoceramus mantelli Merc. ’,
it is difficult to interpret this generic name as anything but a
nomen nudum, because no description or differentiating
characters were given. Inoceramus mantelli de Mercy, 1877,
is based on a number of hinge and outer shell fragments of
large inoceramids from northern France, ranging in age from
Turonian to Coniacian and possibly into the Lower Santo-
nian. Many of these fragments are similar to bilaterally
symmetrical species of immense proportions, not uncommon
in the Chalk of England and northern France, but are
otherwise not particularly different from /noceramus cuvieri
Parkinson, 1819. On this basis it does not therefore seem
necessary to distinguish Platyceramus from Inoceramus sensu
stricto.

In Seitz’s (1961: 54) more valid use of Platyceramus (as a
subgenus of /noceramus), the type species Inoceramus man-
telli de Mercy, 1877, was interpreted according to Barrois’
(1879) description of the species (which was based on ?Conia-
cian specimens). Barrois’ interpretation of J. mantelli differs
from /. cuvieri in having a very much wider umbonal angle, in
which respect it resembles the South African Coniacian
species ‘I’ expansus Baily. It will be necessary to apply to the
I.C.Z.N. if this interpretation of Platyceramus and its type
species is to be accepted.

Seitz had a wide interpretation of the genus and included
many flattish, bilaterally symmetrical inoceramids from vary-
ing parts of the Upper Cretaceous, eg. Inoceramus cycloides
Wegner. Similar large, often smoothish, narrow species also
occur in the Campanian and Maastrichtian. It is difficult to
believe that all these flattish species are correctly assigned to
the same clade and some will need different generic names.
No attempt to solve this taxonomic problem is attempted
here.

Cf. Platyceramus sp. indet.

MATERIAL. A single fragmentary specimen of an almost
smooth, very large species was observed in the main gastro-
pod bed at the base of the Simsima Formation at Jebel Bu

N.J. MORRIS

Milh, section 2. It showed no other characters. The specimen
was not collected, but it indicates the presence of *Platycera-
mus’. This genus is common in the Lower Maastrichtian part
of the St. Lucia Formation, South Africa, and although less
common in the Upper Maastrichtian part of that formation,
the highest Inoceramus horizon seems to consist entirely of
fragments belonging to ‘Platyceramus’.

REFERENCES

Barrois, C. 1879. Sur quelques espéces nouvelles ou peu connues du terrain
crétacé du Nord de la France. Annales de la Societe géologique du Nord,
Lille, 6: 449-S7, pls 4, 5, 12.

Blank, M.R., Krimholtz, L.R., Naidin, D,C. & Sabiuskaya, O.V. 1974. Atlas of
the Upper Cretaceous Fauna of the Don Basin. Ministerstvo Vischego 1
Srednego Spechialinogo OBrazovanii USSR, Communarskii Gornometal-
lurgiieskii Institut, Isdatelestvo N.E.D.R.A. Moscow.

Boehm, J. 1907. Uber Haenleinia nov. subgen. Zeitschrift der Deutschen
geologischen Gesellschaft, Monatsberichte, 59: 317.

— 1909. Uber Inoceramus cripsi auctorum. In Schroeder, H. & Boehm, J.,
1909. Geologie und Palaontologie der subhercynen Kreidemulde. Abhand-
lungen der Koniglich Preussischen geologische Landesanstalt, Berlin, N.F.
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Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):267-273

Issued 30 November 1995

Late Campanian-Maastrichtian Bryozoa from
the United Arab Emirates-Oman border

region

PAUL D. TAYLOR

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD

Synopsis. Seven species of bryozoans have been found encrusting cobbles from the Qahlah Formation (Upper
Campanian or Maastrichtian) of Jebel Huwayyah, east of Al Ain, UAE. This preliminary report describes and
figures the Jebel Huwayyah bryozoan fauna and draws attention to its importance as a rare example of a bryozoan
fauna from the Cretaceous tropics. The following taxa are described: Bullaconopeum nodosum gen. et sp. nov.,
Biaviculigera sp., Pelmatopora sp., Leptocheilopora sp., Tecatia sp., Voigtopora sp. and ‘Berenicea’ sp.

INTRODUCTION

Bryozoans are sessile, suspension-feeding, colonial inverte-
brates, predominantly marine and typically with fossilizable
skeletons of calcite. They often form a major component of
Upper Cretaceous marine biotas: in north-west Europe indi-
vidual assemblages sometimes contain a hundred or more
species (e.g. Voigt 1973; Taylor 1987). Cretaceous bryozoans
from other parts of the world are much less well-known and
appear to be relatively low in both diversity and abundance.
Nevertheless, Upper Cretaceous faunas have been found in
regions as far apart as North America, India and Western
Australia. However, bryozoans of this age have yet to be
described from the Arabian Peninsula.

This paper is a preliminary description of a well-preserved
fauna of encrusting bryozoans from the Qahlah Formation
(Nolan et al. 1990; Skelton et al. 1990) of the Oman Moun-
tains. The Qahlah Formation is late Campanian or early
Maastrichtian in age (A. B. Smith et al, this volume). In the
Jebel Huwayyah area it is thought to represent a transgressive
fan delta depositional system (Skelton ef al. 1990). The
bryozoans were collected from Bed 7 of A. B. Smith et al.
(this volume, Fig. 8) where they encrust cobbles together
with the oyster Acutostrea and colonial corals. Although
Smith ef al. interpreted this bed as a shoreface facies, the
moderately high diversity and general composition of the
bryozoan fauna (with only one malacostegan) suggests a
subtidal origin.

The seven bryozoan taxa found in the Qahlah Formation
extend the geographical ranges of some genera formerly
unrecorded in the Middle East. The bryozoans have particu-
lar significance in providing a rare glimpse of a tropical
Cretaceous bryozoan fauna which can be compared with the
temperate to subtropical faunas found elsewhere, notably in
north-west Europe. Six of the seven species are identified to
genus level only, pending further study and availability of
additional material. The seventh species is distinctive and is
represented by sufficient material to warrant its description as
a new genus and species.

SYSTEMATIC PALAEONTOLOGY

All of the material is from the Qahlah Formation (Upper
Campanian or Maastrichtian), Bed 7, SE corner of Jebel
Huwayyah 1 (see A. B. Smith ef al., this volume), east of Al
Ain, UAE/Oman borders (Sheet NG—40-14D; 1:100,000 grid
reference CM 842878). Specimens are registered in the
collections of The Natural History Museum, London. Scan-
ning electron micrographs were prepared from uncoated
specimens, imaged using back-scattered electrons (see Taylor
1986).

Cheilostome classification used here follows Gordon
(1989), although it is recognized that analysis of phylogenetic
relationships must precede a more satisfactory scheme.

Order CHEILOSTOMIDA Busk, 1852
Suborder MALACOSTEGINA Levinsen, 1902
Superfamily MEMBRANIPOROIDEA Busk, 1854
Family ELECTRIDAE Stach, 1937
Genus BULLACONOPEUM nov.

TYPE SPECIES. Bullaconopeum nodosum sp. nov.; Upper
Cretaceous, Qahlah Formation, Oman Mountains.

NAME. Combination of bulla (Latin, knob, in reference to
the tubercles) with the established bryozoan genus
Conopeum (Latin, mosquito net).

DIAGNOSIS. Colony encrusting, multiserial; autozooidal ope-
sia ovoidal, distal part smooth and crescent-shaped; crypto-
cyst pustulose, broad proximally, moderately broad laterally
but absent distally; gymnocyst narrow, occasionally more
extensive proximally; four gymnocystal tubercles present,
two located near proximo-lateral corners of cryptocyst, two at
distal edges of cryptocyst where crescent-shaped distal part of
opesia begins; kenozooids occasionally developed, irregular
in shape, with pustulose cryptocyst surrounding entire opesia,
lacking the crescent-shaped smooth distal area of autozooids;
pore chambers apparently lacking; intramural buds present,
closure plates not observed; ovicells and avicularia not seen,
presumed absent; early astogeny unknown.

268

REMARKS. This new monospecific genus is distinguished by
the presence of four blunt tubercles around the edges of the
autozooidal cryptocyst. The crescentic shape of the distalmost
part of the opesia arching between the two disto-lateral
tubercles is also characteristic and is presumably more-or-less
coincident with the location of the orifice and operculum in
the living zooid. The absence of ovicells supports assignment
of Bullaconopeum to the malacostegans, a paraphyletic grade
of primitive cheilostomes which at the present day have
non-brooded planktotrophic larvae (cyphonautes). This is
corroborated by the fact that avicularia are also lacking, even
in areas of zooidal crowding where they might be expected to
be found but where kenozooids are developed instead.
Among malacostegan genera, Bullaconopeum resembles
Eokotosokum Taylor & Cuffey, 1992, from the Maastrichtian
of Alberta. This Canadian genus shows a similar develop-
ment of the cryptocyst but has two large disto-lateral spine
bases and no tubercles. Several other malacostegan genera
(e.g. Charixa, Spinicharixa, Villacharixa; see Taylor &
Cuffey 1992) possess spine bases which are lacking in Bulla-
conopeum, although it is possible that the tubercles of
Bullaconopeum represent an evolutionary development from
articulated spines. Tubercles occur in some malacostegan and
other cheilostome genera. Voigt (1992) observed that they
were especially common in bryozoans living in high energy
environments (e.g. on algal fronds), were often variably
present, and suggested that they may function in preventing
mechanical damage. In contrast, the tubercles of Bulla-
conopeum are always present and occur in colonies living in
protected microhabitats such as burrows.

Two families of malacostegans are recognized: Electridae
and Membraniporidae. Bullaconopeum is _ provisionally
assigned to the former family, which is much more diverse,
but confident placement must await the discovery of early
astogenetic stages to ascertain whether the ancestrula is
simple (Electridae) or twinned (Membraniporidae).

Bullaconopeum nodosum sp. nov. Pl. 1
HOLOTYPE. BMNH BZ 3167.
PARATYPES. BMHN BZ 3168 (1), 3172 (1), 3173.

NAME. From nodus (Latin, knot), in reference to the node-
like tubercles.

DESCRIPTION. Colonies are encrusting and sheet-like (Pl. 1,
fig. 1), attaining a maximum diameter of at least 25 mm.
None of the available specimens show unequivocal early
astogenetic stages and therefore nothing is known of the
ancestrula or early budding pattern. Abraded parts of colo-
nies (Pl. 1, fig. 6) show that the basal wall of the zooids are
fully calcified but that pore chambers are lacking.
Autozooids have a rounded rhomboidal shape and are
relatively broad (Pl. 1, figs 2-3). In the holotype frontal
length of the autozooids averages 0-38 mm (range =
0-32-0-47 mm; n = 15) and frontal width 0-35 mm (range =

PLATE 1

P.D. TAYLOR

0-27-0-39 mm; n = 15). An elongate ovoidal opesia occupies |
much of the frontal area of the autozooid. Opesial length in
the holotype averages 0-27 m (range = 0-24-0-30 mm; n =
15) and opesial width 0-19 mm (range = 0-17-0-21 mm; n =
15). Cryptocyst borders the opesia proximally and laterally,
whereas gymnocyst, slightly raised to form a distinctive
semi-circular crescent, defines the distalmost edge of the
opesia. The cryptocyst is pustulose, and is widest and shelf-
like proximally but narrows and slopes more steeply inwards
along the lateral edges of the opesia. The perimeter of the
cryptocyst is slightly depressed relative to the surrounding
gymnocyst, and the cryptocyst:gymnocyst boundary is well-
defined and minutely beaded. The smoothly calcified gymno-
cyst forming the outer edge of the autozooid is narrow,
especially laterally. Four conspicuous but short tubercles
occur along the inner edge of the gymnocyst. Tubercle
diameter is about 40-50 ym. The proximo-lateral pair of
tubercles are located approximately level with, or a little
proximally of, the proximal edge of the opesia. They mark
the transition from the proximal gymnocyst to the narrower
lateral gymnocyst. The disto-lateral pair of tubercles are
situated where the pustulose cryptocyst ends and is replaced
by gymnocyst forming the characteristic crescent-shaped dis-
tal edge of the opesia. Closure plates have not been observed,
although some zooids are plugged with sediment which bears
a superficial resemblance to a closure plate. Intramural buds
are developed in some autozooids as an additional rim of
cryptocyst within the opesia (PI. 1, fig. 5).

Kenozooids are present sporadically (Pl. 1, fig. 4). They
are slightly smaller than the autozooids and more irregularly
shaped. Tubercles are lacking, the opesia is reduced relative
to that of an autozooid, and cryptocyst surrounds the entire
opesia, unlike autozooids where it is absent distally.

Ovicells and avicularia have not been found and are
presumed to be absent.

REMARKS. Bullaconopeum nodosum is the most conspicuous
and commonest of the bryozoans encrusting the collection of
cobbles from Jebel Huwayyah. Colonies occur within pre-
cementation burrows as well as on cobble surfaces where they
are less well-preserved. Colony-form and zooid dimensions
resemble those found in Biaviculigera sp. This similarity is
readily appreciated as superficial when the two species are
studied using SEM (compare PI. 1, fig. 2 with Pl. 2, fig. 1).

Suborder NEOCHEILOSTOMINA d’Hondt, 1985
Infraorder PPEUDOMALACOSTEGOMORPHA d Hondt,
1977
Family CALLOPORIDAE Norman, 1903
Genus BIAVICULIGERA Voigt, 1989

Biaviculigera sp. Pl. 2, figs

MATERIAL. BZ 3171.

REMARKS. A well-preserved specimen of an unidentified

Figs 1-6 Bullaconopeum nodosum gen. et sp. nov., Qahlah Formation (Upper Campanian or Maastrichtian), Bed 7 of Smith et al. (this
volume), SE corner of Jebel Huwayyah 1, east of Al Ain, UAE/Oman borders. 1-4, BMNH BZ 3167, holotype; 1, multiserial arrangement
of zooids, x 35; 2, group of autozooids showing tubercles and pustulose cryptocysts, x 90; 3, autozooid with sediment-filled opesiae, Xx
150; 4, kenozooid partly filled by sediment, x 120. 5-6, BMNH BZ 3168 (1); 5, autozooid (? astogenetically early) at preserved lateral edge
of colony showing proximo-lateral pore window (right of centre) and intramural bud, x 150; 6, abraded growing edge exposing calcified
basal walls of zooids and vertical walls seemingly lacking pore chambers, x 90.

aie

a rs

;

269

LATE CAMPANIAN-MAASTRICHTIAN BRYOZOA

270

species of Biaviculigera has a sheet-like encrusting colony
with spineless autozooids, non-pustulose cryptocysts, avicu-
laria and ovicells. This genus (type species Membranipora
praecipua Brydone, 1914) was proposed by Voigt (1989) for
nine species of Cenomanian-Maastrichtian cheilostomes,
seven of which were formerly assigned to the genus Membra-
nipora as used in its broadest sense. Biaviculigera is charac-
terized by having two types of avicularia. In the Qahlah
Formation species these comprise large avicularia with spatu-
late rostra and small avicularia with pointed rostra.

Suborder ASCHOPHORINA Levinsen, 1909
Infraorder CRIBRIOMORPHA Lang, 1916
Family PELMATOPORIDAE Lang, 1916
Genus PELMATOPORA Lang, 1916

Pelmatopora sp. Pie 2F hiee2

MATERIAL. BZ 3170.

REMARKS. Pelmatopora is a diverse genus which ranges from
the Coniacian to the Upper Campanian or possibly Danian
(Larwood 1962). The Qahlah Formation species has about
16-20 arched costae, each with a large pelma (pore) in the
narrow median area of costal fusion. Pelmatidia and lateral
costal fusions seem to be absent, and ‘interzooecial secondary
tissue’ (sensu Larwood 1962) and oral spines have not been
observed. However, sediment obscures much of the morpho-
logical detail. A few zooids bear ovicells, and many have
small paired avicularia located on either side of the high
autozooidal orifice, inwardly directed towards the orifice,
spatulate and with well-developed transverse bars.

Genus LEPTOCHEILOPORA Lang, 1916

Leptocheilopora sp. RIS ation 3

MATERIAL. BZ 3169 (3).

REMARKS. An inconspicuous colony of Leptocheilopora
encrusts one of the cobbles. The small autozooids have about
14-16 closely-spaced costae forming a flat frontal shield. Four
oral spines are present and there are numerous small avicu-
laria, each with a relatively large proximal gymnocyst and a
spatulate rostrum without a transverse bar. Lateral costal
fusions and pelmata have not been observed. The orifice has
well marked lateral constrictions, dividing a deep poster from
the distal anter. None of the species of Leptocheilopora
revised by Larwood (1962) compares closely with the Qahlah
Formation species, especially with regard to the avicularia
which are uncommon in this genus. In Europe Lep-
tocheilopora ranges from the Santonian to the Maastrichtian
(Larwood 1962).

PLATE 2

P.D. TAYLOR

Infraorder HIPPOTHOOMORPHA Gordon, 1989
Family HIPPOTHOIDAE Fischer, 1866
Genus TECATIA Morris, 1980

Tecatia sp. Ple2, figs

MATERIAL. BZ 3168 (2).

REMARKS. The tiny autozooids of this ascophoran are dis-
persed widely across the substratum, and arranged in a
seemingly chaotic manner. It is unclear whether the wide-
spacing of the autozooids is because they have long caudae,
are interspersed with stolon-like heterozooids, or reflects loss
of intervening zooids. The autozooidal orifice has a distinct
sinus, and the strongly arched gymnocystal frontal wall shows
a line of pore windows just above substratum level. Although
this species is assigned to Tecatia, the minute pores in the
frontal wall which are characteristic of Tecatia have not been
observed, possibly because of preservational limitations.

Tecatia is represented by four species from the Upper
Maastrichtian of the type area and one Recent species from
the Pacific coast of North and Central America (Morris 1980;
Voigt & Hillmer 1983). Voigt (1987) described how T. minuta
Morris, 1980, can be found in ‘minicaverns’ formed by
thalassinoid burrows in hardgrounds in the Maastrichtian
Chalk Tuff. Similarly, the Qahlah Formation species of
Tecatia inhabited the cryptic habitats provided by precemen-
tational burrow systems in cobbles.

Order CYCLOSTOMATA Busk, 1852
Suborder TUBULIPORINA Milne Edwards, 1838
Family STOMATOPORIDAE Pergens & Meunier, 1886
Genus VOIGTOPORA Bassler, 1952

Vc. ztopora sp. Pl. 2, figae

MATERIAL. BZ 3169 (1), 3172 (2).

REMARKS. A few colonies of the runner-like cyclostome
Voigtopora encrust the cobbles. This genus is distinguished
from the related Stomatopora by the presence of lateral
branch ramifications and the long proximal extensions of the
zooids which flank the previous zooid in linear series (see
Illies 1976). Voigtopora appears to range from the Hauteriv-
ian to the Campanian in Europe (Pitt & Taylor 1990), and
also occurs in the Albian and Cenomanian of Texas. The
species-level systematics of this genus is in need of revision,
with characters of early colony development likely to be of
considerable importance.

Figs 1-6 Bryozoans from the Qahlah Formation (Upper Campanian or Maastrichtian), Bed 7 of Smith et al. (this volume), SE corner of
Jebel Huwayyah 1, east of Al Ain, UAE/Oman borders; 1, Biaviculigera sp., BMNH BZ 3171, part of colony showing autozooids with
smooth cryptocysts, a large spatulate avicularium (in recess, centre left), and several small acuminate avicularia, x 60; 2, Pelmatopora sp.,
BMNH BZ 3170, autozooid (partly obscured by sediment) showing large pelmata in median area of costal fusion, x 100; 3,
Leptocheilopora sp., BMNH BZ 3169 (3), autozooid flanked by two small, abraded avicularia, x 230; 4, Tecatia sp., BMNH BZ 3168 (2),
oblique view of a zooid with sediment-plugged orifice (arrow) and line of basal pore windows, x 250; 5, Voigtopora sp., BMNH BZ 3169
(1), branch with three zooids viewed obliquely, x 45; 6, ‘Berenicea’ sp., BMNH BZ 3169 (2), part of colony showing transversely-wrinkled

frontal walls, x 45.

LATE CAMPANIAN-MAASTRICHTIAN BRYOZOA

271

Family incertae sedis
Genus ‘BERENICEA’ Lamouroux, 1821

‘Berenicea’ sp. Pl. 2, fig. 6

MATERIAL. BZ 3169 (2).

REMARKS. Part of a sheet-like tubuliporine cyclostome is
present at the edge of one of the burrows. The zooidal frontal
walls are transversely wrinkled, but in the absence of gono-
zooids it is impossible to assign the colony to a genus.
Therefore, following the procedure adopted by Taylor &
Sequieros (1982) and Pitt & Taylor (1990), the specimen is
placed in the informal genus “Berenicea’.

DISCUSSION

This new bryozoan fauna has both palaeobiogeographical and
palaeoecological importance. As noted above, the great
majority of Cretaceous bryozoan faunas have been described
from Europe. Nothing was previously known of Upper
Cretaceous bryozoan faunas from the Arabian Peninsula.
The Qahlah Formation specimens therefore fill a major
geographical gap in our knowledge of Upper Cretaceous
bryozoans. Furthermore, as the late Cretaceous proto-Oman
Mountains would have been located almost on the palaeo-
equator (e.g. A. G. Smith ef al. 1994), it provides a tropical
fauna that can be compared with the temperate faunas of
Europe and elsewhere. Six of the seven genera found in the
Qahlah Formation occur also in north-west Europe, the
exception being Bullaconopeum. At the genus-level, there-
fore, it is seems that there was little taxonomic differentiation
between late Cretaceous temperate and tropical bryofaunas.
In addition, albeit based on a small number of species, the
proportion of cheilostomes (five species) to cyclostomes (two
species) in the tropical Qahlah Formation is fairly typical for
a fauna of this age (cf. Lidgard et al. 1993, fig. 7).

Along with other cemented invertebrates and plants, bryo-
zoans are commonly found encrusting the surfaces of cobbles
like those from the Qahlah Formation. Cobbles provide a
variety of different microhabitats for these sessile organisms,
including upper and lower external surfaces, and crevices
formed by vacated borings or pre-cementational burrows.
Therefore, cobbles allow the study of small scale niche
differentiation among demonstrably in-situ organisms. Few
studies of cobble-dwellers have been undertaken in the fossil
record, and the only detailed study involving a diverse
Cretaceous bryozoan fauna is that of Wilson (1986) on
cobbles from the Aptian Faringdon Sponge Gravel of
England. Wilson found a clear distinction between species
colonizing the outer surfaces of the cobbles, which were
robust in morphology, and species living in vacated borings,
which often had a delicate construction. This he interpreted
as a consequence of the physical rigours experienced by the
encrusters on outer surfaces compared with the protected
burrows. Preliminary study suggests that a similar differentia-
tion may occur among the colonizers of the Qahlah Forma-
tion cobbles, although further field sampling and careful
mapping of distributions will be needed to substantiate this
impression. It would be particularly instructive to compare
the cyclostome-dominated Aptian Faringdon Sponge Gravel

P.D. TAYLOR

cobbles with the cheilostome-dominated younger Qahlah
Formation cobbles.

ACKNOWLEDGEMENTS. For discussion and advice on matters bryozoo-
logical and stratigraphical, I wish to thank Ehrhard Voigt, Peter
Skelton, Noel Morris, Andrew Smith and Andrew Gale.

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LATE CAMPANIAN-MAASTRICHTIAN BRYOZOA

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Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):275-276

Issued 30 November 1995

Maastrichtian brachiopods from the United
Arab Emirates-Oman border region

ELLIS F. OWEN

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD

INTRODUCTION

Brachiopods were particularly rare in the Maastrichtian sedi-
ments of the United Arab Emirates-Oman borders region.
Despite intensive collecting only four specimens were found,
all coming from the lower Loftusia-rich beds of the Qahlah
Formation exposed at Jebel Huwayyah (see introduction for
locality details). These four specimens belong to a hitherto
undescribed genus of terebratulid.

SYSTEMATIC DESCRIPTIONS

Class TEREBRATULIDA Waagen, 1883
Suborder TEREBRATULIDINA Waagen, 1883
Superfamily TEREBRATULACEA Gray, 1840

Family TEREBRATULIDAE Gray, 1840

Genus PSEUDOGIBBITHYRIS nov.

TYPE SPECIES. Pseudogibbithyris arabica sp. nov.

DIAGNOsIS. Medium-sized uniplicate terebratulid, slightly
longer than wide. elongate-oval in general outline and evenly
biconvex. Umbo short, beak suberect, foraman small, perme-
sothyrid. Cardinal process present, brachial loop simple.

REMARKS. Pseudogibbithyris differs from Gibbithyris and
Concinnithyris from the European Upper Chalk (Senonian),
which it resembles in external morphological features, in its

Fig. 1. Pseudogibbithyris arabica gen. et sp. nov., holotype,
BMNH BF47; Jebel Huwayyah, section 2, beds 3-6. A, dorsal; B,
lateral; C, anterior views: all x 1.

distinctly flat bifid cardinal process, deep hinge-trough and
short, triangular hinge-plates.

Pseudogibbithyris arabica sp. nov. Figs 1, 2

DIAGNOsIs. As for genus.
Types. Holotype, BMNH BF47; paratypes BMNH BF4446.

DESCRIPTION. The dorsal valve is dominated by a low
median fold bounded by faint carinae originating from a point
midway between the umbonal area of the valve and the
anterior margin of the shell. A corresponding shallow sulcus
in the ventral valve forms a wide, shallow uniplication
occupying the whole width of the anterior commissure.
Internal characters include a well-developed flat, but dis-
tinctly bifid cardinal process, and a deep hinge-trough. The
hinge-plates are short, triangular in outline with a slightly
concave ventral surface, and are deflected towards the dorsal
valve. The triangular shape of the hinge-plates is maintained
in the development of the horizontally placed bands of the
descending branches of the brachial loop, which is uncompli-
cated and terminates in a very high arcuate transverse band.

REMARKS. The only morphological features which this spe-
cies has in common with other Terebratulidae of the Upper
Cretaceous are the distinctly oval general outline and unipli-
cate anterior commissure. The internal structure, notably the
distinctive cardinalia and short triangular hinge-plates, are
features more typical of terebratulid species of the Upper
Jurassic to Lower Cretaceous, the closest comparison being
to species of Nucleata and Pygites. However, it is not sug-
gested that there is any direct relationship between the
species described here as Pseudogibbithyris arabica gen. et
sp. noy. and the Tithonian genera mentioned above. As no
reference to any forms which could be confidently compared
to this species are known, it is treated as a previously
undescribed genus and species.

It would be unwise at this stage to draw any firm conclu-
sions about the phylogenetic relations of this taxon. The
simple cardinalia and brachial loop structure seen in the
transverse serial sections (Fig. 2), is unusual and suggests a
late Jurassic or early Cretaceous ancestry. However, the
taxon should remain broadly assigned until more material is
obtained, allowing further investigation.

OCCURRENCE. All specimens came from the Loftusia-rich
facies (beds 3-6) Jebel Huwayyah, section 2 (see volume
Introduction for locality details). They are Maastrichtian in
age.

276 E.F. OWEN

Fig. 2. Sixteen transverse serial sections through the holotype of Pseudogibbithyris arabica gen. et sp. nov. (BMNH BF47) from Jebel
Huwayyah, section 2, beds 3-6. Note the bifid cardinal process (sections 5 & 6), the short triangular hinge-plates, and high, arcuate
transverse band of the brachial loop. The numerals denote the distance in mm between each section. All x 3.

Bull. nat. Hist. Mus. Lond. (Geol.) 51(2):277-305

Issued 30 November 1995

Late Campanian-Maastrichtian rudists from
the United Arab Emirates-Oman border

region

N. J. MORRIS

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD

P.W. SKELTON

Department of Earth Sciences, The Open University, Walton Hall, Milton Keynes, Buckinghamshire

MK76AA

CONTENTS

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References

Se cee ea ee GME Oe See ic eld oleae ctu elas a cluninis wis ec derteloster ogee 277
Pomp Geb aT RRR aDcOGe ae cOc oD ean eben cncascce qa concaC cnn eMeRseHonn tn acts 278
ap sjopebec candle soi aeaale deta sie face eee erie ea ee nteeteaaae a seats 303

Synopsis. About 30 species of Upper Campanian and Maastrichtian rudist bivalves from the Qahlah and Simsima
Formations of the United Arab Emirates-Oman border region are described. They include Semailia smithi gen. et
sp. nov. of the Dictyoptychidae, Glabrobournonia arabica gen. et sp. nov. of the Radiolitidae (Biradiolitinae), and
Pseudosabinia gen nov. of the Radiolitidae (Joufiinae). The geological succession at Qarn Mulayh (Mileih), 7 km

west of Jebel Buhays, is described.

STRATIGRAPHY AND AGE

In addition to the stratigraphy outlined by Smith, Morris and
Gale at the beginning of this volume we include here strati-
graphical details of the successions at Qarn Mulayh and Qarn
Murrah (the ‘red qarns’) situated to the west of Jebels Buhays
and Faiyah, UAE (Smith ef al., this volume, fig. 4). At these
localities, limestone platform sedimentation on the serpenti-
nized front of the Semail Ophiolite started earlier than in the
jebels to the east, and is equivalent in age to the arenaceous
facies of the Qahlah Formation.

A section at Qarn Mulayh is given in Fig. 1. Similar beds
also form the majority of the outcrop at Qarn Murrah, to the
north. On the basis of the rudists, we suspect that the
succession at both qarns is of late Campanian age, and may
possibly extend into the earliest Maastrichtian. These lime-
stones correlate laterally with the sandy facies of the Qahlah
Formation, with only a few of the species present above in the
basal beds of the Simsima Formation.

Table 1 gives a full list of the rudists collected, and their
distribution is shown in Fig. 2. The oldest fauna (1 on Fig. 2)
occurs at Qarn Mulayh and Qarn Murrah, and also in the
lower Qahlah Formation gravels at Jebels Huwayyah and Bu
Milh. It is probably Campanian in age (the M1 fauna of
Skelton et al., 1990). The middle fauna (2 on Fig. 2) occurs in
the upper Qahlah Formation Loftusia-Beds at Jebel
Huwayyah, and the top Qahlah Formation gravels (with
acteonellids) at Jebel Bu Milh. It is Campanian/early Maas-
trichtian in age (the intermediate M1/M2 fauna of Skelton et

al., 1990). The upper fauna (3 on Fig. 3) occurs in the main
part of the Simsima Formation, at Jebels Faiyah, Buhays,
Rawdah and Huwayyah. It is Maastrichtian in age.

The level 1 fauna compares closely with the so-called
Pironaea-Pseudopolyconites Senonian fauna of Sladic¢-
Trifunovic (1989), for which the type area is the “Vrbovac
Beds’ of eastern Serbia. Diagnostic novel taxa (not present in
the older Gosau Beds) are: *Pironaea, Yvaniella,
*Pseudopolyconites, Joufia, Neoradiolites, Branislavia
(?=*Colveraia), Sabinia |= * Pseudosabinia here), and Mitro-
caprina (asterisks denote those also in the UAR/Oman
fauna). In addition, the following were noted by Sladic-
Trifunovié: *Vaccinites loftusi, V. ultimus, V. orientalis, V.
bacevicensis, Hippurites colliciatus, ?*H. lapeirousei, * Hippu-
ritella cornucopiae. Moreover, Sladic¢-Trifunovic (1989: 149)
observed: ‘The species *Vaccinites oppeli Douvillé, also
found in the Vrbovaé Reef (Baéeviéa), is certainly of a special
biostratigraphic importance, since it was previously believed
to exist only in Early Senonian’.

Sladié-Trifunovié (1989: 153), following the earlier works
of Milovanovic and Grubi¢, regarded the assemblage as
Upper Campanian/Maastrichtian, mainly based on the sup-
posed evolution of Pironaea. She did, however, acknowledge
the arguments of others (for an earlier-extending range),
based on orbitoids, and agreed that the fauna was probably
absent from the Upper Maastrichtian. Pejovi¢ & Radoicic
(1987) for example, had revised the age of the ‘Bra¢ Marbles’
(with Pironaea etc) to the Lower-Middle Campanian. Sladic¢-
Trifunovié (1989: 154) responded: ‘If the ‘Bra¢ Marbles’,
which include P. milovanovici [Kitihn’s species, regarded as

278

Table 1 Systematic list of the rudists found in the Qahlah and
Simsima Formations.

Family CAPROTINIDAE Gray, 1848
Genus GYROPLEURA Douvillé, 1887
Gyropleura sp.

Family PLAGIOPTYCHIDAE MacGillavry, 1937
Genus PLAGIOPTYCHUS Matheron, 1843
Plagioptychus cf. toucasianus Matheron, 1843

Family DICTYOPTYCHIDAE Skelton in Skelton and Benton, 1993
Genus DICTYOPTYCHUS Douvillé, 1905
Dictyoptychus morgani (Douvillé, 1904)
Genus EODICTYOPTYCHUS Skelton & El-Asa’ad, 1992
Eodictyoptychus aff. arumaensis Skelton & El-Asa’ad, 1992
Genus SEMAILIA Morris & Skelton, gen. nov.
Semailia smithi Morris & Skelton sp. nov.
Semailia sp.

Family HIPPURITIDAE Gray, 1848
Subfamily TORREITINAE Grubic, 1979

Genus Torreites Palmer, 1933

Torreites sanchezi (Douvillé, 1927) milovanovici Grubic, 1980
Subfamily HIPPURITINAE Gray, 1848

Genus VACCINITES Fischer, 1887
Vaccinites loftusi (Woodward, 1855)
Vaccinites vesiculosus (Woodward, 1855)
Vaccinites oppeli (Douvillé, 1892)

Genus HIPPURITES Lamarck, 1801
Hippurites aff. lapeirousei Goldfuss, 1841
Hippurites cornucopiae Defrance, 1821
Hippurites aff. cornucopiae Defrance, 1821

Genus PIRONAEA Meneghini in Pirona, 1868
Pironaea cf. polystyla Pirona, 1868

Family RADIOLITIDAE d’Orbigny, 1847
Subfamily RADIOLITINAE d’Orbigny, 1847
Genus Praeradiolites Douvillé, 1902
Praeradiolites cf. subtoucasi Toucas, 1907
Genus Radiolites Lamarck, 1801
?Radiolites sp.
Subfamily PPEUDOPOLYCONITINAE Sladi¢-Trifunovic, 1983
Genus Pseudopolyconites Milovanovi¢, 1937
Pseudopolyconites aff. parvus Milovanovic, 1935
Subfamily BIRADIOLITINAE Douvillé, 1902
Genus Biradiolites d Orbigny, 1850
Biradiolites aff. baylei Toucas, 1909
? Biradiolites aff. baylei Toucas, 1909
Genus Glabrobournonia Morris & Skelton gen. nov.
Glabrobournonia arabica Morris & Skelton sp. nov.
Sub-family SAUVAGESIINAE Douvillé, 1908
Genus Durania Douvillé, 1908
Durania cf. gaensis (Dacqué, 1903)
Durania cf. apula (Parona, 1900)
Durania form A
Durania form B
Durania spp.
Subfamily LAPEIROUSIINAE Kihn, 1932
Genus Lapeirousia Bayle, 1878
Lapeirousia sp.
Genus Osculigera Kiihn, 1932
Osculigera cf. vautrinioides Vogel, 1970
Subfamily JOUFIINAE Karacabey-Oztemiir 1981
Genus Colveraia Klinghardt, 1921
Colveraia aff. variabilis Klinghardt, 1921
Genus Pseudosabinia Morris & Skelton gen. nov.
Pseudosabinia aff. klinghardti (Boehm, 1927)

N.J. MORRIS AND P.W. SKELTON

‘advanced’], were accepted as being of the Upper Campanian
age, there would be no sense in talking about the evolution of
the genus Pironaea’. The Pironaea story has, indeed, been
shown to be incorrect by J. M. Pons & E. Vicens (unpub-
lished), who suggested that the different forms reflect ontog-
eny, rather than phylogeny, while Swinburne (1990: 27)
dated the Bra¢ Marbles (= Pucis¢a Formation) as early-
middle Campanian, based on Sr isotope correlations, in
agreement with Pejovic & Radoicic. Moreover, by the same
means, she and others (Swinburne et al., 1992) re-assigned
some Pironaea beds in Bulgaria, which had previously been
placed at various levels in the Maastrichtian, to the Campa-
nian and lowest Maastrichtian. Thus the ‘Pironaea-
Pseudopolyconites Senonian’ fauna seems to have ranged
from the early Campanian to the earliest Maastrichtian.
Additionally, Philip & Platel (1987) assigned their Torreites
beds in Dhofar to the Campanian, based on orbitoids, while
in the Caribbean the range of Torreites sanchezi (in our level
1 fauna) is also restricted to the Campanian (Rojas, Iturralde-
Vinent, & Skelton, in press).

In summary, a Campanian age for the level 1 fauna seems
the most plausible (though on the rudist evidence alone, we
still cannot yet exclude the earliest Maastrichtian). On the
other hand, the level 3 fauna presents a marked contrast,
having only V. oppeli, H. cornucopiae, Colveraia and Pseu-
dosabinia in common with the classic Pironaea-
Pseudopolyconites Senonian fauna. H. cornucopiae, though,
is also well known from the Maastrichtian of Sicily and
elsewhere. As noted in Skelton et al. (1990: 545), this
younger fauna is also characterized by some distinctive
Arabian/Iranian endemics (e.g. Dictyoptychus). The Level 3
fauna is independently dated by ammonites (Kennedy, this
volume) as late Early to early Late Maastrichtian.

SYSTEMATIC PALAEONTOLOGY

Most of the described and figured rudists are in the collec-
tions of The Natural History Museum, and are cited with the
prefix BM; many of them have the additional prefix LL, and
unless otherwise stated the specimens are in the Morris, Gale
& Smith Collection; most of the other material is in the
Skelton collection, collected by Skelton and Nolan (Skelton
et al, 1990). The morphotype nomenclature used, eg. ‘eleva-
tor’, ‘clinger’, ‘recumbent’, is according to the scheme of Gili
& Skelton (1994), summarised by Skelton (1991) and Ross &
Skelton (1993).

Family CAPROTINIDAE Gray, 1848.
(emend. Skelton, 1978; =Monopleuridae Munier-Chalmas,
1873)
Genus GYROPLEURA Douvillé, 1887

TYPE SPECIES. Requienia cenomanensis d’Orbigny, 1850.

REMARKS. There are no accessory cavities.

Pl. 1, fig. 2

MATERIAL. Four small specimens from top bed 4 or basal
bed 5, Simsima Formation Jebel Rawdah, section 1, ammo-
nite and inoceramid horizon; two are attached to the upper
valve of an ‘Umbonium’, BM LL41767-69.

Gyropleura sp.

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS

Metres

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Comments

- Some 30 m, exposed only sporadically
through dunes: red-stained bioclastic
packstones to wackestones, with large
gastropods (Campanile), irregular
echinoids and small radiolitid rudists.

- Localized clusters of moderate to
large-sized elevator to radial clinger
rudists, in life position: Pironaea,
Vaccinites, Torreites, Osculigera.
Shelly packstone matrix.

- Bioclastic packstone with scattered
flakes of radiolitid left valves.

- Dense radiolitid flake floatstone.
- Packstone with Vaccinites and rare

Torreites. Basal lag with Pseudosabinia.

- Biostrome of Osculigera and others.

- Fine grainstone to packstone, with
drifts of radiolitid flakes, and isolated
Pironaea, Colveraia and Pseudosabinia.

- Grainstone with Pseudosabinia.

- Grainstone with toppled Pironaea and
Torreites, and radiolitid floatstone cap.
- Nodular shelly packstone with
burrowed top. Rare small radiolitids,
burrowing bivalves and Cyclolites.

- (sand-covered)

- Bouquets of small radiolitids

- Sugary dolomitised calcarenite with
limonitic (?ex-serpentinite) grains.
Scattered Osculigera, small radiolitids,
Plicatula, cyclolitid corals and regular
echinoids. Omphalocyclus present.

- (some 20 m sand-covered)

- Coarse serpentinite and limestone
clast conglomerate.

Deepening

?Beach

Shallow carbonate bank with mobile lime sands and rudist meadows

Interpretation

KEY

Bioclastic limestone

Shell-rich bioclastic
limestone

Orbitoid-rich bioclastic
limestone

Conglomerate

Elevator rudist
Recumbent rudist

Radiolitid flakes
Cyclolitid coral

Burrowed surface

Serpentinite pebble

279

‘Fig. 1 Measured section at Qarn Mulayh (Mileih), 7 km west of Jebel Buhays; section logged at the north end of the western flank by P W

| Skelton.

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N.J. MORRIS AND P.W. SKELTON

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2 e e e e @
] e e ? e e«© ee @ e e @ e

Fig. 2. Stratigraphical distribution of the rudists. Ut = stratigraphical units: 1 = lower part of the Qahlah Formation, 2 = upper part of the
Qahlah Formation (the gravels and the Loftusia-Beds), 3 = Simsima Formation. Ps = Pseudopolyconites, Bi = Biradiolites, Bo =
Glabrobournonia, Du = Durania, La = Lapeirousia, Os = Osculigera, Pk = Pseudosabinia aff. klinghardti, Pr = P. ‘rtanjica’, Co =
Colveraia, Eo = Eodictyoptychus, Se = Semailia, Di = Dictyoptychus, P| = Plagioptychus, Pi = Pironaea, V\ = Vaccinites loftusi, Vv =
Vaccinites vesiculosus, Vo = Vaccinites oppeli, Hc = Hippurites cornucopiae, H = Hippurites aff. lapeirousei, T = Torreites.

DESCRIPTION. All four specimens have the valves conjoined,
with the upper left valve slightly exogyriform and the lower
valve rather longer but with considerable variation in size of
attachment surface; line of commissure slightly sinuous.
Outer shell surface of both valves with fine, evenly and
closely spaced, radiating striiform ribs. Shell margins crenu-
late. Adductor attachments visible in the left valve of speci-
men LL41767 on concave surfaces of the hinge plate, a little
below the plane of commissure. Gyropleura is a clinger.

Family PLAGIOPTYCHIDAE MacGillavry, 1937: 105, 152
(ex. Plagioptychinae MacGillavry, 1937 (plagioptychinés
Douvillé, 1888: 729))
Genus PLAGIOPTYCHUS Matheron, 1843.

TYPE SPECIES. Plagioptychus paradoxus Matheron, 1843,
subsequently designated by Kutassy (1934: 172).

REMARKS. Species of Plagioptychus are clingers to low eleva-
tors.

Plagioptychus cf. toucasianus Matheron, 1843 PI. 1, fig.
1

cf. 1843 Plagioptychus toucasianus Matheron, 117.

MATERIAL. Two small specimens from near the base of the
Simsima Formation at Jebel Faiyah, BM LL41765 (centre),
LL41766 (from section 2), Skelton Collection.

DESCRIPTION. Inequivalve, right valve smaller than free left
valve, exogyriform with sinuous commissural margin with
large attachment area, surface slightly rugose with prominent
commarginal growth lines. The rather short exterior dorsal
surface has a sub-vertical ligament groove and the shell
surface bulges to the posterior of this line. Outer calcitic shell
layer thin to medium, with thin inner layer without pallial
canals. Prominent tooth and attached myophore projects into
upper valve.

Free left, upper valve globose, regularly coiled, gryphaeate
with smooth surface; outer calcitic shell layer thin, inner layer
recrystallized to calcite but medium to thick, with radially
elongated narrow canals that do not pass into polygonal

PLATE 1

Fig. 1 Plagioptychus cf. toucasianus Matheron, from Jebel Faiyah, lower part of Simsima Formation. 1a, anterior view, 1b, section through

structure and in that way differ from Mitrocaprina or the
upper valve of Coralliochama. The narrow canals also do not
bulge internally as is common in some forms of the genus.
Anterior myophore robust and nearly level with the commis-
sure. An oblique septum runs from the anterior tooth to the
ventral margin. The cavity posterior to this septum houses the
large central tooth and conjoined posterior myophore in a
dorsal position.

Family DICTYOPTYCHIDAE Skelton in Skelton & Benton,
1993
(ex. Trechmannellidae Cox, 1933: 65)

DIAGNOSIS. Inequivalved rudists, attached by right valve.
Valves uncoiled and ligament absent. Outer (calcitic) shell |
layer compact. Inner (originally aragonitic) shell canaliculate
throughout, in both valves. Two projecting teeth in left valve,
straddling ridge-like central tooth in right valve. Posterior
tooth dorso-ventrally flattened, flanking body-cavity, and
separated from the dorsal margin by a small accessory cavity,
which may be a relic of the ligamentary cavity. Anterior
myophoral platforms extended both dorsally on hinge plate,
around anterior tooth, and ventrally from hinge plate. Poste-
rior myophore of left valve projecting, with adductor scar
facing outwards, into recess, or socket, in posterior wall of
right valve, and directly flanking body-cavity.

REMARKS. The distinctive features of this family were dis-
cussed, in relation to Eodictyoptychus, by Skelton and
El-Asa’ad (1992). The dictyoptychid myocardinal apparatus
differs from that seen in the Caprinidae (s.s.), the Plagiopty-
chidae, and Sabinia (s.s.) in all of which the posterior tooth
and posterior myophore (with inward-facing muscle scar) of
the left valve are separated from the body-cavity by the
combined central tooth socket and an accessory cavity,)
extending ventrally from it, which receives the salient myo-
phore of the right valve. The latter cavity is itself demarcated
by a prominent lamina running from the anterior tooth to the
postero-ventral margin of the valve. No equivalents of this)
lamina, and the associated accessory cavity, are present in the)
left valve of dictyoptychids.

Pseudosabinia (Radiolitidae) differs in its possession of a

left valve approximately 5 mm from the commissure, BM LL41766, x 1, Skelton Collection 1c, dorsal view, BM LL41765, x 1.

Fig. 2 Gyropleura sp., from Jebel Rawdah, section 1, loose from top of bed 2 or bed 3 of Simsima Formation; 1a, posterior view, 1b, dorsal
view, Ic, view looking down on left valve, BM LL41768, x 2.

Fig. 3 Dictyoptychus morgani (Douvillé), from Jebel Buhays, section 1, bed 11 of Simsima Formation, side view, BM LL41680, x 0-5.

Fig. 4 Eodictyoptychus arumaensis Skelton & Al-Asa’ad, from Jebel bu Milh, section 1, top of Qahlah or basal Simsima Formation; 4a, top
view, 4b, internal view, 4c, dorsal view, BM LL41927, left valve, x 1-5.

281

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS

282
ligamentary invagination, and the presence of cellulopris-
matic structure in the outer shell layer of the right valve.
Despite the similarity to the Antillocaprinidae MacGillavry,
1937, in their lack of a ligament, and the presence of the small
dorsal accessory cavity, which may be a relic of it, the
dictyoptychids differ from Antillocaprina (at least) in having
the posterior myophore of the left valve projecting, rather
than parallel with the plane of commissure. Moreover, the
outer shell layer of antillocaprinids is invariably very thin,
whilst that of dictyoptychids is usually relatively thick in the
right valve.

Genus DICTYOPTYCHUS Douvillé, 1905.
(nom. nov. for Polyptychus Douvillé 1904, non Huebner,
1816)

TYPE SPECIES. Polyptychus morgani Douvillé, 1904, by
monotypy.

SYNONYM. Trechmannella Cox, 1933 (obj.).

REMARKS. Attached right valve with large polygonal canali-
cular structure. Outer layer of right valve thicker than that of
left valve, with exposed growth margin; sharply peaked
ridges on right valve growth margin in some specimens. We
agree with Pons et al., 1992, who regard all described species
as probably synonymous.

Dictyoptychus morgani (Douvillé, 1904) Pipe ties'3

1904 Polyptychus morgani Douvillé: 248-51, pl. 33 bis.
1905 Dictyoptychus morgani (Douvillé); Douvillé: 198.

MATERIAL. Jebel Faiyah, section 1, bed 3, BM LL41670-71;
section 2, LL41659; Jebel Faiyah, centre, LL41683-—84. Jebel
Agabah, Simsima Formation, loose, BM LL41676. Jebel
Thanais, Simsima Formation, loose in scree, BM
LL41668-69. Jebel Buhays, section 1, bed 11, LL41677,
41680; section 1b, LL41674; section 1a, Simsima Formation,
BM LL41675. Jebel Rawdah, section 1, bed 3, BM
LL41657-58, 41679, 41681; section 2, bed 19, LL41673;
section 2, loose on scree, LL41667; section 4, bed 1,
LL41672; Jebel Rawdah, north, LL41690. Jebel Bu Mihl,
section 1, bed 3, BM LL41660-66. Jebel Huwayyah, section
2, beds 3-5, BM LL41678; Jebel Huwayyah, LL41685-89.
Jebel Sa’ah, basal Simsima Formation, BM LL41682.

REMARKS. Occurs widely in the lower half of the Simsima
Formation, sometimes in great numbers. Dictyoptychus is a
multigeniculate elevator (see Ross & Skelton, 1993: fig. 5.2).

Genus EODICTYOPTYCHUS Skelton & El-Asa’ad, 1992

TYPE SPECIES. Eodictyoptychus arumaensis Skelton &

El-Asa’ad, 1992, by original designation.

PLATE 2

N.J. MORRIS AND P.W. SKELTON

Eodictyoptychus aff. arumaensis Skelton & El-Asa’ad,
1992. Pl. 1; fig. 4; Pl 25 fig: 2

vy aff. 1992 Eodictyoptychus arumaensis Skelton &

El-Asa’ad: 108-13, pl. 1, figs 1-6.

MATERIAL. Jebel Bu Milh, section 1, Qahlah/Simsima For-
mation Boundary, BM LL41927-8; section 2, LL41929; all
are left valves. A possible fragment of a right valve from
Qarn Murrah, BM LL41977, Skelton Collection.

DESCRIPTION. Moderately inaequivalved, free left valve with
thin outer calcitic shell layer, inner aragonitic layer recrystal-
lised but formed of varyingly sized canals of polygonal
section, even within the teeth and the massive myophores.
Two subequal, projecting teeth in free valve, the anterior one
below the umbo, the smaller, posterior one ventral to this,
the two separated by a socket whose centre lies above and
towards the umbo from the posterior tooth. The teeth lie on
the body-cavity side of a massive plate which includes the
attachment surfaces of the myophores, and takes up more
than half of the area within the valve when looking down on
the commissural plane.

We have not been able to detect the accessory cavity dorsal
to the posterior tooth, but this may be due to the difficulty of
its preparation. The projecting posterior myophore is sepa-
rated from the body-cavity by a narrow ledge which is not
present in the type material from Khashm Buwaibiyat,
approximately 50 km north of Riyadh.

REMARKS. Eodictyoptychus varies from a lateral clinger to a
recumbent.

Genus SEMAILIA Morris & Skelton, gen. nov.

TYPE SPECIES. Semailia smithi Morris & Skelton gen. et sp.
nov.

DIAGNOSIS. Bicornate and multicarinate, subequivalve with
a thin outer shell layer in each valve. Inner shell canaliculate
in both valves. Myocardinal arrangement typical of dictyopty-
chids with two teeth in the left valve, the posterior tooth
dorso-ventrally flattened and flanking the body-cavity with-
out an intervening accessory cavity. No ligament present. The
posterior myophore of the left valve projecting into a socket
in the right valve, again flanking the body-cavity without an
intervening accessory cavity.

REMARKS. The absence of a posterior accessory cavity in the
left valve excludes this taxon from the Caprinidae, Plagiopty-
chidae and Sabinia s.s., and the outward facing, ie monopleu-
riform, myophores, are typical of the dichtyoptychid plan.
Unlike other members of the Dictyoptychidae this genus does
not show the differentially greater thickening of the outer
calcitic layer of the right valve.

Fig. 1 Semailia smithi Morris & Skelton, gen. & sp. nov. Jebel Huwayyah, section 2, Loftusia-Beds; 1a, anterior view, 1b, posterior view,

Ic, view of right valve, Id, dorsal view; holotype, BM LL41931, x 1.

Fig. 2 Eodictyoptychus aff. arumaensis Skelton & Al-Asa’ad, Jebel bu Milh, section 1, top of Qahlah or base of Simsima Formation; internal

view of left valve, BM LL41928, x 1.

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS

283

284

N.J. MORRIS AND P.W. SKELTON

Fig. 3 Semailia smithi Morris & Skelton sp. nov. Four sections through the holotype approximately 10 mm either side of the commissure,
BM LL41931; a, b, right valve, MC — accessory cavity for posterior myophore of left valve, PO — posterior tooth socket, AC — accessory
cavity dorsal to tooth socket, CI — central tooth, AO — anterior tooth socket; ¢, d, left valve, with fine outer and large irregularly polygonal

inner canals; X 1.

PLATE 3

Fig. 1 Semailia sp., Jebel bu Milh,section 2, base of Simsima Formation; 1a, ventral view of broken right valve, 1b, dorsal view of right

valve; BM LL41932 x 0-75.

Fig. 2 Torreites sanchezi milovanovici Grubi¢; Haushi-Hugf Massif, Eastern Oman, BM LL41975, Samir Hanna Collection; 2a, view looking
down onto commissural plane with broken left valve partly preserved in situ, X 1; 2b, dorsal view showing intucking of pillar L, 2c,

posterior view showing infolds of pillars, 2d, ventral view; 2b-d, x 0-5.

Fig. 3 Vaccinites aff. oppeli (Douvillé), Jebel bu Milh, section 2, Simsima Formation, bed 10, BM LL41730, marginal surface of outer shell

layer of right valve, x 1.

Semailia smithi Morris & Skelton sp. nov. PI. 2, fig. 1,
Fig. 3

MATERIAL. A single specimen, the holotype, BM LL41931,
from Jebel Huwayyah, section 2, Loftusia-Beds.

DIAGNOSIS. Typical of the genus but with three strong cari-
nae or flaring radial costae on each valve.

DESCRIPTION. Both valves are preserved, closed, although
the body-cavity is matrix-filled and includes larger foramin-
ifera. The specimen has two serpulids attached, one to each
valve and aligned radially with respect to the umbones, with
their apertures close to the valve commissure, amid the plicae
of the ventral margins.

The shell is curvingly biconical, with the two valves sepa-
rately following almost a semicircular direction of growth, in

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 285

286

a single plane, so that the umbones or apices are beginning to
approach each other; sub-equivalve with the right valve
slightly ‘longer’ than the left. Three prominent carinae or
flaring radial costae, skewed to the posterior, two at approxi-
mately 180°, set anteriorly and posteriorly on both valves and
the third normal to these on the ventral margin. Dorsal
margin rounded. Shell surface otherwise relatively smooth on
both valves.

No ligament present. Very thin outer calcitic shell layer in
both valves. Canals are present in the thick inner shell layer
of both valves, round and capillary-like around the margins,
becoming larger inside, irregularly rounded polygonal in
section, especially in the left valve.

Two teeth in the left valve are typical of the family, the
anterior one is rounded and knob-like, the posterior one is
dorso-ventrally flattened. One central tooth in the right
valve. The posterior myophore of the left valve projects into
an accessory cavity (socket) in the right valve. The anterior
myophore of the left valve is a broad shelf and has large
canals. There is an accessory cavity in the right valve which
lies dorsally to the flattened posterior tooth socket.

COMMENTS. The crescentic form of the shell suggests a
recumbent life position, possibly reclining on the dorsal
flank. It was found in a matrix of marly limestones with large
specimens of Loftusia with a similar matrix filling the body-
chamber, forming a loftusid packstone.

COMPARISON WITH OTHER TAXA. There is a great similarity
in the myocardinal arrangement with that of other dictyopty-
ichids, but Semailia differs in having a much thinner outer
shell layer in both valves and having strong shell carinae.

Pl. 3, fig. 1

MATERIAL. A single specimen from Jebel Bu Milh, section 2,
basal Simsima Formation, BM LL41932.

Semailia sp.

DESCRIPTION. The specimen is a large right valve, its anterior
and posterior part nearly symmetrical about a dorso-ventral
plane. Multicarinate but otherwise smooth. Two wide ante-
rior and posterior carinae form a wide kite-shape in dorsal
view. Dorsal surface flattish near the commissure, but
umbones incurved and separating two gently concave ante-
rior and posterior areas with a low mid-dorsal carina that is
prominent at the umbo, but faces towards the mid-dorsal
margin. Ventral part of shell has three sub-equal strong
carinae, which are equidistant from the dorsal margin and
inclined towards the anterior.

Outer shell layer thin and now formed of structureless
recrystallized blocky calcite. Thick canaliculate inner shell
layer with capillary-like polygonal canals throughout, includ-
ing the teeth, fine at the margins becoming larger inwards,
now also recrystallized to calcite.

Right valve has part of projecting stout central tooth
preserved behind base of the socket for the anterior tooth of

PLATE 4
Fig. 1

N.J. MORRIS AND P.W. SKELTON

the left valve and in front of the socket of the posterior tooth
of the left valve, containing a fragment of that dorso-ventrally
flattened tooth.

COMPARISON WITH OTHER SPECIES. Semailia sp. differs from
S. smithi in having two additional ventral carinae. There is
not enough material to know if this difference is significant in
discriminating separate species.

Family HIPPURITIDAE Gray, 1848
Subfamily TORREITINAE Grubic, 1980
Genus TORREITES Palmer, 1933

TYPE SPECIES. Hippurites (Vaccinites) sanchezi Douvillé,
1927, by original designation.

DESCRIPTION. Outer shell layer of right valve with tight
marginal infoldings, giving rise to radiating crests (Douvillé,
1894) around the shell margin of the right valve, but variable
in extent and number. Outer shell layer of left valve thin,
postero-dorsally digitiform, smooth except for fine growth
lines on the upper surface, but becoming overgrown with
epibionts radiating from its low apex. Inner shell layer of both
valves partly canaliculate, canals of right valve relatively
large, sub-radial and sub-rectangular in section, in the area of
the anterior muscle attachment. We have observed a similar
reticulate network of vertical ridges separating tabulate
canals in the anterior myophoral ledge of Vaccinites gosavien-
sis (BM 33972, a specimen with the original aragonite pre-
served). The canals of the left valve are narrow and radiate
from the apex.

REMARKS. Philip & Platel (1994) pointed to the similarity of
Torreites with their new genus, Praetorreites, from the Lower
Campanian, Samhan Formation of south-east Oman. The
latter has canaliculate structure of the inner shell layer of the
left valve and regular pedunculate radial structures in the
outer shell layer of the right valve, which, they claim, are
comparable with the radiating marginal crests of Torreites.
They compare the subfamily, raised to family rank, with both
the Plagioptychidae and the Hippuritidae, concluding that
the similarity is greater with the Plagioptychidae. If Philip &
Platel are correct in their suggestion that the Torreitinae are
not hippuritids, then it follows that the three large infolds of
the outer shell layer of the right valve are not homologues of
the hippuritid pillars. The origin of two of the pillars of
Torreites is claimed to be from the two small ‘pillars’ of
Praetorreites and considered by Philip & Platel to be ana-
logues of pillars ‘E’ and ‘S’. Philip & Platel’s plate 8, fig. 1
shows the position of these ‘pillars’ in relation to the teeth
and myophores of the upper valve. If this arrangement is
compared with the hinge structures of Dictyoptychus (Dou-
villé, 1904: pl. 33 bis, fig. 4), the alignment of the sockets and
teeth is very similar, although they are at a more obtuse angle

Torreites sanchezi milovanovici Grubi¢, Haushi-Hugf Massif, Eastern Oman (also figured Skelton & Wright, 1987: pl. 67, fig. 1), BM

LL28004, view looking down onto right valve (note coarse cellular structure of myophore in lower part of figure and tooth sockets to the

right of pillar ‘L’), x 1.

Fig. 2 Vaccinites aff. oppeli (Douvillé), Jebel bu Milh, section 2, Simsima Formation limestone, BM LL41733, view of naturally broken

section of right valve, x 0-4.

Fig. 3 Vaccinites vesiculosus (Woodward), Jebel Huwayyah, Loftusia-Beds, BM LL41716, Skelton Collection, 3a, view of partly eroded left

valve, 3b, view of right valve, x 0-5.

renee eee aaa iii eee inated aeeeeiena eked piaiinas = eeeaieaaedin aiTiaieaan aioe Ela EtEaiSeEe aE ceREGE eA enE — —

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 287

288 N.J. MORRIS AND P.W. SKELTON

to the ventral shell margin. This suggests a relationship
between Dictyoptychus and Praetorreites.

It is difficult to relate the anterior infolding of Torreites to
the position of ligamentary invagination of the plagiopty-
chids, but much easier if Torreites is interpreted as a hippu-
ritid. Details of the morphology are illustrated by Skelton &
Wright (1987: fig. 2). In Skelton & Wright’s interpretation of
Torreites loss of the hippuritid canal system of the outer shell
layer of the left valve is by recurvature of the shell margin to
expose the mantle margins. The umbo of the operculiform
left valve is close to the ventral margin.

Philip & Platel (1994: fig. 4) claim a diphyletic origin for
the ‘three-pillared’ genus Torreites and thus claim to have
refuted the palaeobiogeographical interpretation of Skelton
& Wright (1987). Their diagram shows Praetorreites occur-
ring before Torreites and giving rise independently to the
Caribbean and Arabian species. However, their date for the
small Caribbean species Torreites tschoppi on this diagram
does not show its full range. There is ample evidence to show
that this species is best dated as Santonian in Cuba (Rojas et
al., in press) and therefore pre-dates Campanian Praetorre-
ites. Their analysis leading to the view that the torreitines are

not closely related to the hippuritids relies heavily on the Fig. 4 Torreites sanchezi milovanovici Grubié; smooth top surface
interpretation of the ancestral status of Praetorreites and the of left valve with advancing front of epibionts (arrowed), BM
supposed homology of the pedunculate folds, with the mar- LL41975, x 2.

ginal radiating crests of Torreites, which in the latter are a
consequence of tight infoldings of the outer shell surface. A
superficially similar structure in section may be seen in the
outer shell layer of the lower valve of Dictyoptychus striatus,
but this is formed as a consequence simply of salient radial
ridges on the growth margin of that shell layer, without any
infolding of the outer surface. In Philip & Platel’s (1994: pl. 7,
fig. 3) illustration of Praetorreites the similarity, if anything,
seems greater with the radial ridges of the dictyophychid,
than with the intuckings of Torreites. In all particulars Prae-
torreites is similar to Eodictyoptychus and appears to have
little in common with Torreites.

The criticism of Skelton and Wright’s (1987) explanation of
the distribution of Torreites is therefore unconvincing. We Fig. 5 Camera lucida drawing of Vaccinites loftusi (Woodward);
maintain the view that the differences between the Caribbean from Qarn Murrah, BM LL41933, Skelton Collection, x 1.
and Arabian Torreites do not warrant greater that sub-

specific separation, and that their similarities do reflect that Torreites sanchezi (Douvillé 1927) subsp
genetic interchange between the two areas of occurrence did OVA O MEAG Enbic 1980 ; F

take place. ; ;
a Pl. 3, fig. 2; Pl. 4 fig. 1
1927 Hippurites (Vaccinites) sanchezi Douvillé: 54, 55, pl. 4,
fig. 1.
1980 Torreites milovanovici Grubic¢: 92, 93, pl. 1, fig. 1, fig.
4,
PLATE 5

Figs 1,2 Hippurites aff. lapeirousei Goldfuss; 1, Jebel Faiyah, section 1, Simsima Formation, bed 6, BM LL41754, mass of variously
orientated individuals, x 0-5; 2, Jebel Faiyah, section 1b, Simsima Formation, bed 2, BM LL41755, view looking down onto a pair of right
valves, X 2.

Fig. 3 Vaccinites vesiculosus (Woodward), Jebel Huwayyah, Loftusia-Beds, BM LL41973, V. Chalmers Collection, view looking down on
eroded left valve exposing pillars of right valve, x 1.

Figs 4-7 Hippurites cornucopiae Defrance; 4-6, Jebel Faiyah, loose from low in the Simsima Formation, Skelton Collection; 4, BM
LL41747, view of left valve (pores appear polygonal and radially vermiculiform where less eroded), x 1; 5, BM LL41744, eroded left valve
showing pores and canals, x 1; 6, BM LL41745, view of left valve with oscules and pores appearing vermiculiform or polygonal depending
on the degree of erosion, x 1; 7, BM LL41737, Jebel Thanais, natural section through right valve showing slightly pedunculate pillars, x 1.

Fig. 8 Hippurites aff. cornucopiae Defrance, Jebel Rawdah, southern flank, loose from Simsima Formation, BM LL41753, Skelton
Collection, natural section of right valve, x 0-5.

Fig. 9 Pironaea cf. polystyla Pirona, Qarn Mulayh, BM LL41938, Skelton Collection, section through right valve, x 1.

289

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS

290

1994 Torreites milovanovici Grubi¢; Philip & Platel: pl. 8,
fig. 4, fig. 4.
(For further synonomy see Skelton & Wright, 1987)

TYPE SPECIMEN. The holotype of the subspecies is BM
LL27699, Iraq Petroleum Company Collection (CP 86), on
weathered surface of section, 7-7 m from top at Qarn
Mulayh.

OTHER MATERIAL. Two specimens from the same horizon on
Qarn Murrah, BM LL41761, 62; a well-preserved specimen
showing details of the upper valve, from the Haushi-Huqf
Massif, eastern Oman, S. Hanna Collection, BM LL41975;
the material of Skelton & Wright (1987) from eastern Oman,
BM LL28004; one specimen from Jebel Huwayyah, section 1,
Qahlah gravels, BM LL42763; Qarn Mulayh, no specimens
collected.

COMMENTS. BM LL41975 shows a boundary between shiny
outer shell material and encrusted surface on the outer
surface of the upper valve which we interpret as representing
the limit of encroachment upon the exposed left mantle
surface by epibionts. BM LL28004 shows the loose canalicu-
lar structure of the anterior muscle attachment area of the
right valve together with the tooth sockets.

Subfamily HIPPURITINAE Gray, 1848
Genus VACCINITES Fischer, 1887

TYPE SPECIES. Hippurites cornuvaccinum Bronn, 1931, by
monotypy.
Vaccinites loftusi (Woodward, 1855)

1855
1897

Fig. 5

Hippurites loftusi Woodward: 58, pl. 3 figs 2-3.
Hippurites loftusi Woodward; Douvillé: 210, pl. 33
(17), figs 1, la, 1b.

Vaccinites loftusi (Woodward); Toucas: 82, 83, figs
WA, WF.

1904

MATERIAL. Three specimens from Qarn Murrah, BM
LL41933, LL41969-70; another from Jebel Huwayyah,
Loftusia-Beds, LL41934; Skelton Collection.

REMARKS. These specimens have the typical coarse external
ribs of Woodward’s species but are very poorly preserved
internally. The pillars of a Qarn Murrah specimen resemble
those of Woodward’s type but were not enhanced by section-
ing. The pores of the upper valve are in the form of coarse
polygons and the outer surface of this left valve has radial
undulations. Woodward’s type material has incipient multiple
infoldings of the outer shell layer of the right valve, remins-
cent of Pironaea.

PLATE 6
Fig. 1
concave smooth dorsal radial band to the right, x 1.
Fig 2
ventral view, 2c, anterior view; all Xx 1-5.

N.J. MORRIS AND P.W. SKELTON

Vaccinites vesiculosus (Woodward, 1855) PI. 4, fig. 3;
Pl. Satiens

Hippurites vesiculosus Woodward: 59, pl. 4, fig. 6.
Hippurites vesiculosus Woodward; Douvillé: 201, pl.
29 (13), figs 6, 7.

Vaccinites vesiculosus (Woodward); Toucas: 110, 111.

1855
1897

1904

MATERIAL. Qarn Murrah, BM LL41691-7, LL41721-26,
Skelton Collection; Jebel Huwayyah, section 1, bed 9, BM
LL41698-702; section 2, bed 7 (the main coral bed), BM
LL41703+4; section 2, beds 2-7, BM LL41705-15 plus
another 9 specimens; Jebel Huwayyah, loose, Skelton Collec-
tion, BM LL41716—-20, V. Chalmers Collection, BM
LL41973; three possible specimens from Jebel Thanais, loose
in scree below lowest 3m of the Simsima Formation, BM
LL41935-37.

REMARKS. The specimens from the Loftusia-Beds at Jebel
Huwayyah are larger than either those from Qarn Murrah or
Woodward’s type material. They however have the same type
of pillars in the right valve with a long thin aréte cardinal and
pedunculate pillars. The complex pattern of the canals and
pores of the left valve are also similar in each of the three
groups of material. V. vesiculosus is the dominant rudist
species in the Loftusia-Beds at Jebel Huwayyah.

Vaccinites aff. oppeli (Douvillé, 1892) Pl. 3, fig. 3; Pl. 4

igs 2

aff. 1866 Hippurites dilatatus Defrance; Zittel: 142, pl. 24,
figs 1-5.

aff. 1881 Hippurites zitteli Munier-Chalmas in Zittel: 83,
fig. 118 (non Matheron, 1880).

aff. 1892 Hippurites oppeli Douvillé: 36, 37, figs 23, 24, pl.
4 (18), fig. 5.

aff. 1897 Hippurites oppeli Douvillé: 203, pl. 31 (15), figs 1,
la.

aff. 1904 Vaccinites oppeli (Douvillé); Toucas: 109, 110, pl.
17 (18), figs 2, 2a.

MATERIAL. Jebel Bu Mihl, section 2, common, many may be
in life position, lower part of the main Simsima Limestone,
BM LL41728-30, 41733; Hill to south of Jebel As-Saifr (east
of Jebel Huwayyah), BM LL41731-32, Skelton Collection.

REMARKS. In this very large form, the pores of the upper
valve are present in one specimen and are similar to the pore
system in topotypic material from Gosau, particularly on the
outer third of the radius. The outer shell surface is rather
smooth. The raised pattern on the growth surface of the outer
shell layer of the right valve is extremely similar to the
Austrian material (cf. Zittel, 1866: fig. 1). The pillars in the
right valves of our material match very closely with those of

Durania Form A, Jebel Huwayyah, section 1, from the Qahlah gravels, BM LL41948, part of outer shell layer of right valve with

?Radiolites sp., Jebel bu Milh, section 2, beds 7/8 at the Qahlah/Simsima boundary, BM LL41947, right valve, 2a, view of interior, 2b,

Figs 3,4 Praeradiolites cf. subtoucasi Toucas, Jebel Rawdah, section 4, bed 1, basal rudist bed of Simsima Formation; 3, BM LL41941,
section across lower part of right valve, x 1; 4, BM LL41942, view of anterior of right valve, x 1.

Fig. 5 Pseudopolyconites aff. parvus Milovanovi¢, Qarn Mulayh, BM LL41974, Skelton Collection, x 1; 5a, section through right valve with
aréte cardinale at top; 5b, surface of part of right valve with smooth ventral radial band centre and sediment with spines to right.

Fig. 6 Durania cf. apula Parona, Jebel Rawdah, section 2, Simsima Formation, bed 10, BM LL41951; 6a, ventral view of right valve showing
radial bands; 6b, part of upper surface of right valve with small holes near the radial bands, x 1.

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS 291

J ‘
yo

292

the Austrian material. The differences seem to relate only to
phyletic size increase.

Genus HIPPURITES Lamarck, 1801

TYPE SPECIES. Hippurites bioculatus Lamarck, 1801, by
monotypy.

Hippurites aff. lapeirousei Goldfuss, 1841 PI. 5, figs 1, 2

aff. 1895 Hippurites lapeirousei Goldfuss; Douvillé: 164, pl.
24 (11), figs 7-10.

Orbignya lapeirousei Goldfuss; Toucas: 53, pl. 6
(12), figs 10, 11.

aff. 1903

MATERIAL. Jebel Faiyah, Sharjah, section la, bed 6, BM
LL41754; section 1b, bed 2b, BM LL41755; Jebel Faiyah,
loose, BM LL41758, Skelton Collection; Jebel Rumaylah,
BM LL41756, Skelton Collection; Jebel Mundassah, BM
LL41757, Skelton Collection.

REMARKS. This species is present in large masses of disorien-
tated specimens close to coral patches, low in the Simsima
Formation at Jebel Faiyah. It has broad and short pillars and
lacks an aréte cardinale. It differs from typical specimens
from Maastricht in having a rather smooth shell surface to its
right valve.

Pl. 5, figs 4-7

aff. 1821 Hippurites cornucopiae Defrance: 195, pl. 58, figs
la, 1b (only).
1910 Hippurites (Hippuritella) cornucopiae Defrance;
Douvillé: 79, pl. 7, figs 3-5.

MATERIAL. Jebel Faiyah, section 1, BM LL41735; Jebel
Faiyah, loose, BM LL41738, 4174447, Skelton Collection;
Jebel Buhays, section 1, bed 15, BM LL41739-43; section 3,
BM LL41736; Jebel Thanais, BM LL41737, Skelton Collec-
tion; Jebel Rawdah, section 2, BM LL41734.

Hippurites cornucopiae Defrance, 1821

REMARKS. The pores of the upper valve, together with the
disposition and shape of the pillars of the lower valve, match
those described by Douvillé from Sicily, which is the type
locality of Defrance’s species. However, although the pores
are polygonal and denticulate even when the surface is only
slightly eroded, they are radially vermiculiform when the
outer surface is intact. Hence the species should be assigned
to Hippurites not Hippuritella (pace Douvillé, 1910). Rather
conical specimens occur at the coral patch horizon at Jebel
Faiyah, sometimes within the coral clumps.

Hippurites aff. cornucopiae Defrance, 1821 PI. 5, fig. 8

MATERIAL. Jebel Faiyah, BM LL41750, LL41752, Skelton
Collection; Jebel Rawdah, section 3, above Simsima conglo-
morate, BM LL41748; Jebel Rawdah, loose, BM LL41749,
41751, 41753, Skelton Collection.

PLATE 7

N.J. MORRIS AND P.W. SKELTON

REMARKS. Specimens from Jebel Faiyah and Jebel Rawdah
that occur well up the sequence of the Simsima Formation are
of Upper Maastrichtian age. They are similar in general plan
to H. cornucopiae but are much larger, always being more
than twice the diameter of the latter.

Genus PIRONAEA Meneghini in Pirona, 1868
TYPE SPECIES. Hippurites polystylus Pirona, 1868, p. 511.

Pironaea cf. polystyla Pirona, 1868 Piles) ioe 9.

1868 Hippurites polystylus Pirona: 511.

MATERIAL. Three specimens from Qarn Mulayh, loose from
lower part of sequence, BM LL41938—40, Skelton Collection;
the species was also seen at Qarn Murrah (see Skelton ef al.
1990: fig. 9a).

DESCRIPTION. Outer shell layer of medium thickness with
numerous secondary pillars. Inner shell layer of right valve
slightly thicker than outer shell layer, recrystallised.

COMMENTS. Swinburne et al. (1992) have shown that the
supposed evolutionary sequence of Pironaea ‘species’
described as Maastrichtian in Serbia are more likely to be of
Campanian to earliest Maastrichtian age.

Family RADIOLITIDAE d’Orbigny, 1847
(as Radiolidae, emend. Gray, 1848: 439)
Subfamily RADIOLITINAE d’Orbigny, 1847
Genus PRAERADIOLITES Douvillé, 1902, p. 467

TYPE SPECIES. Radiolites fleuriaui d’Orbigny, 1842, from the
Cenomanian of Le Mans, by original designation.

DESCRIPTION. Ligamentary invagination of right valve usu-
ally present. Right valve elongate, left valve operculiform.

Praeradiolites cf. subtoucasi Toucas, 1907 Pl. 6, figs 3, 4

cf. 1907 Praeradiolites subtoucasi Toucas: 31, pl. 3 (13),
figs 8, 9.

MATERIAL. Four specimens from Jebel Rawdah, section 4,
from the Simsima Formation, bed 1, BM LL41941—-44.

DESCRIPTION. A slightly distorted attached (right) valve,
‘D’-shaped in transverse section, which has developed a
secondary bilateral symmetry about a dorso-ventral plane.
Moderately wide radial bands are separated by a narrower
sinus, on the ‘flat face’ which is formed at the postero-ventral
margin. The two radial bands are wider than the central sinus
and all three are deep below the plane of commissure.
Internally the symmetry differs from the outside, with the
ligamentary invagination set at about 30 degrees to the
posterior of the axis bisecting the external ‘D’ shape.

COMPARISON WITH OTHER SPECIES. The sinuses are deeper

Figs 1-4 Durania cf. gaensis (Dacqué), Jebel bu Milh, section 1, Qahlah Formation; all x 0-75; 1, BM LL41922, individual from first clump,
natural section showing shell layers and tabulae of right valve and part of surface and myophore of left valve; 2, BM LL41923, individual
from second clump, commissural surface of right valve; 3, BM LL41924, individual with domed left valve from third clump, internal mould
with some shell adhering; 3a, postero-dorsal view; 3b, basal view of right valve; 3c, anterior dorsal view; 4, BM LL41922, second individual

from first clump; 4a, postero-dorsal view; 4b, postero-ventral view.

293

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS

294

than those of P. aristidis (Munier-Chalmas, 1888) as figured
by Toucas (1907, pl. 6 (16), figs 9, 10). They are more similar
to P. boucheroni (Bayle) as figured by Toucas (1907, pl. 3
(13), fig. 10 only) and to P. subtoucasi Toucas (1907, pl. 3
(13), figs 8, 9).

Genus RADIOLITES Lamarck, 1801

DESCRIPTION. Widely biconical with upper valve shorter
than lower right valve. Outer layer of fixed right valve thick
with calcite cellular structure radially stretched. Aréte cardi-
nale usually present, usually short.

Pl. 6, fig. 2

MATERIAL. Three specimens From Jebel Bu Milh, section 1,
from the Qahlah/Simsima boundary, BM LL41945—46; Jebel
Bu Milh, section 2, beds 7, 8, BM LL41947; one probable
specimen from Jebel Rawdah, section 2, from scree,
LL41980.

?Radiolites sp.

DESCRIPTION. The simple smooth radial bands, the flaring
growth rugae-like stacked projecting cones, and the radially
aligned cell pattern of the outer shell layer of the right valve
are similar to Radiolites, particularly the groups of Radiolites
radiosus and R. sauvagesi (Toucas 1908). The radial bands
are not thrown into stong folds and therefore this species does
not belong to Praeradiolites. No aréte cardinale is visible and
it may well be absent. This is a common trend in many rudist
lineages and we suspect that it may happen independently in
this form of Radiolites.

Subfamily PPRUDOPOLYCONITINAE Sladi¢-Trifunovic,
1983b, p. 239
(ex. Pseudopolyconitidae Sladi¢-Trifunovic1983b, emend.
herein)

Genus PSEUDOPOLYCONITES Milovanovic, 1937
[Pseudopolyconites Milovanovi¢, 1935, was invalid because
no type species was originally designated, in contravention

of ICZN Rules, Art. 13A (b)]

TYPE SPECIES. Pseudopolyconites parvus Milovanovic¢, 1935.

Pseudopolyconites aff. parvus Milovanovic, 1935 PI. 6,
fig. 5

Pseudopolyconites parvus Milovanovié: 188, 252
(nomen nudum).

Pseudopolyconites parvus Milovanovic: 54-70,
figs 1b-8.

Pseuodopolyconites parvus Milovanovi¢; Milo-
vanovic: 4-14, figs 2-9.

1934
aff. 1935

aff. 1937

MATERIAL. Two specimens from Qarn Murrah, BM
LL41978-79, and one specimen collected loose at Qarn
Mulayh, BM LL41974; all Skelton Collection.

PLATE 8

N.J. MORRIS AND P.W. SKELTON

REMARKS. The specimen LL41974 is well-preserved and
shows the spines and a wide smooth ventral radial band. The
aréte cardinale is long and narrow, with a typically rounded
to ovoid distal end. The left valve LL41978 shows the aréte
cardinale and the spine bases on its upper surface.

Subfamily BIRADIOLITINAE Douvillé, 1902
Genus BIRADIOLITES d@ Orbigny, 1850

TYPE SPECIES. Biradiolites canaliculatus d’Orbigny, 1850
(ICZN Opinion pending).

Biradiolites aff. baylei Toucas, 1909 Pl’ 8; hes 133

aff. 1909 Biradiolites baylei Toucas: 119, pl. 24 (9), figs 9,

10.

) Biradiolites royanus (d’Orbigny); Toucas: 103,

pl. 19 (4), figs 34-38.

Biradiolites aquitanicus Toucas: 107, pl. 20 (5),

fig. 20.

21967 Biradiolites bulgaricus Pamouktchiev: 35, pl. 1,
fig. 4; pl. 2, figs 3, 4.

MATERIAL. Five specimens from Jebel Huwayyah, section 2,
four from the top of the Loftusia-Beds, BM LL41799-41802,
one from the Loftusia-Beds, BM LL41815, Skelton Collec-
tion; one specimen from Jebel Bu Milh, loose, but probably
from the basal gastropod bed of the Simsima Formation at
Jebel Bu Milh, BM LL41805.

1909

cf. 1909

DESCRIPTION. Right valve with secondary approach to bilat-
eral symmetry. Dorsal margin convex-alate, with a central
(dorsal) raised portion, which is itself centrally grooved in a
small specimen. Has strong ‘lateral’ carinae which curve
downwards, giving the appearance of a stretched bow when
the smooth dorsal surface is viewed. Ventral margin of right
valve with wide radial bands separated by an acute carina that
forms a prominent downward ‘V’ at about the same level as
the ‘lateral’ carinae.

REMARKS. B. baylei was a lateral clinger, on its broadly
expanded antero-dorsal face. The latter feature, and the
anterior-ward leaning of the interband, suggest Toucas’
group of B. fissicostatus, of which B. baylei is the Maastrich-
tian representative.

A specimen in the Trechmann collection, identified by
Chubb (1971) as Bournonia thiadensi Vermunt from the
Maastrichtian of Jamaica has similar plications on the ventral
face of the lower valve, but a wider and flatter central area on
the reverse side and in that way resembles the next species.

?Biradiolites aff. baylei Toucas, 1909 Pl. 8, figs 3-5

MATERIAL. Six specimens from the central eastern face of
Jebel Faiyah, loose on lower part of the Simsima Formation,
BM LL41793-98, Skelton Collection; one specimen from
Jebel Rumaylah, from the lower Dictyoptychus level,
LL41814, Skelton Collection; two specimens from Jebel

Figs 1,2 Biradiolites aff. baylei Toucas, Jebel Huwayyah, section 2, Loftusia-Beds, beds 3-8; x 1-5; 1, BM LL41800; 1a, dorsal view; 1b,
view of left valve; le, ventral view; 2, BM LL41799; 2a, view of left valve; 2b, ventral view; 2c, dorsal view; 2d, basal view of right valve.
Figs 3-5 __Biradiolites ?aff. baylei Toucas, Jebel Faiyah centre, loose from lower part of Simsima Formation, Skelton Collection; x 0-75; 3,
BM LL41794, 3a, ventral view; 3b, basal view of right valve; 3c, dorsal view; 4, BM LL41796; 4a, view of left valve; 4b, ventral view; 4c,

dorsal view; 5, BM LL41795; 5a, view of left valve; 5b, ventral view.

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS

295

296

Faiyah, section 1, in hard limestone, ?from the lower Sim-
sima Formation, BM LL41803-04; two specimens from Jebel
Buhays, section 3, lower Simsima Formation, BM
LL41810-11; three specimens (two conjoined) from Jebel
Aqabah, BM LL41812-13; one specimen from Jebel Raw-
dah, section 2, bed 13, BM LL41806; one specimen from the
same section, bed 23, BM LL41807; two others from the
same section, loose, BM LL41808-09; one specimen from
Jebel Rawdah, BM LL41816, loose, Skelton Collection; one
specimen from Jebel Ja’Alan, southern Oman Mountains,
west side, lower Simsima Formation, BM LL41817, Skelton
Collection.

COMMENTS. Similar to Biradiolites aff. baylei but with an
additional carina between the dorsal radial band and the
posterior carina. The radial bands are relatively narrower
than in B. aff. baylei. A number of species of this complex
have been described by Pamouktchiev (1967); Pons et al.
(1992) in a study of specimens from Somalia suggested that
the distinctions are unjustified.

Genus GLABROBOURNONIA Mortis & Skelton gen. nov.

TYPE SPECIES. Glabrobournonia arabica Morris & Skelton
sp. nov.

DIAGNOSIS. Small genus with a cornute lower valve and a
very low, slightly convex upper valve. Right valve smooth
except for fine growth lines and three major sinuses in the
shell margin which leave sinusoidal traces, one at the ventral
margin, one at the dorsal margin and one centrally on the
posterior margin. Upper valve with reticulate sculpture of
fine radiatig ribs and concentric growth laminae.

REMARKS. At present we are aware of only one species.
Glabrobournonia differs from Bournonia in being devoid of
ribbing on the right valve.

Glabrobournonia arabica Morris & Skelton sp. nov.
Plo 9 igs 12

HOLoTyPE. BM LL41873 from Jebel Rawdah, section 1, the
lower Simsima Formation, bed 2, just below low Upper
Maastrichtian ammonites and inoceramids.

PARATYPES. Jebel Rawdah, section 1, bed 2, LL41870-72;
bed 3 and top bed 2, LL41874—95; loose, LL41869; section 4,
bed 2, the basal rudist bed, LL41896-41916; Jebel Rawdah,
southern flank, scree from Lower Simsima Formation,
LL41818-22, Skelton Collection; Jebel Faiyah, section 2,
lower Simsima Formation, LL41855—57; Jebel Thanais, lower
Simsima Formation, LL41854; Jebel Buhays, section 1,
LL41836-51, LL41917-—21; LL41825-27, Skelton Collection;

PLATE 9

N.J. MORRIS AND P.W. SKELTON

section 1, lower Simsima Formation, beds 4-10, LL41832-35;
section 1b, loose, BM LL41852-S3.

OTHER MATERIAL. Qarn Murrah, 50-60 m from bottom of
section, LL41828 (16 specimens), LL41829-31, Skelton Col-
lection; Jebel Huwayyah, section 1, beds 10-11, Loftusia-
Beds, LL41858; section 2, beds 2-7, Loftusia-Beds,
LL41859-68; LL41823-24, Skelton Collection.

DIAGNOSIS. As for genus.

DESCRIPTION. Mostly smooth horn-shaped lower valve with
two bands representing a downward sinuosity where the
cellular structure is exposed; one band is anterior, the other is
ventral, and there is a further posterior sinuosity where
growth increments are more obvious than the cellular struc-
ture. Upper valve has exposed cellular structure radiating
from excentric umbo, otherwise smooth, gently convex with a
sinuous margin to fit lower valve.

Subfamily SAUVAGESIINAE Douvillé, 1908
Genus DURANIA Douvillé, 1908

TYPE SPECIES. Hippurites cornupastoris Des Moulins, 1826,
from the Turonian of France.

REMARKS. The specimens of Durania from the Qahlah For-
mation of Jebel Bu Milh are well-preserved, but even this
material does not give us sufficient information concerning
ecophenotypic variation. Different ‘morphs’ from different
horizons are listed separately but we do not know whether
they are different species.

Durania cf. gaensis (Dacqué, 1903) Pl. 7, figs 14

cf. 1903 Radiolites ga’ensis Dacqué: 374, pl. 35, figs 7-9.
cf. 1909 Sauvagesia gaensis (Dacqué); Toucas: 85, pl. 16
(17), figs 3-5.
21909 Sauvagesia flicki Toucas: 84, 85, pl. 16 (17), figs
6-8.
cf. 1910 Durania gaensis (Dacqué); Douvillé: 50.

LOCALITIES OF PREVIOUSLY FIGURED MATERIAL. Dacqué’s
type material came from Ga’a near Abu Roash, Egypt, and
was said by Douvillé (1910) to be Turonian in age; Toucas’
material occurs with Lapeirousia and was said to occur from
Coniacian to Maastrichtian in Tunisia.

MATERIAL. Three clumps from Jebel Bu Milh, section 1,
from the upper part of the Qahlah Formation, LL41922-24;
two further doubtful clumps from Jebel Faiyah, one loose
from section 1, the second from section 1b, bed 2b, BM
LL41925-26.

DESCRIPTION. Rather large, outer shell layer of nght valve

Figs 1,2 Glabrobournonia arabica Morris and Skelton gen. nov., sp. nov.; 1, Jebel Rawdah, section 2, Simsima Formation, bed 2, BM
LL41873, holotype; la, ventral view; 1b, posterior view; 1c, dorsal view; 1d, anterior view; X 1-5; 2, south end of Jebel Buhays, lower part
of Simsima Formation, BM LL41825, paratype, Skelton Collection; view of internal mould of left valve; x 1-5.

Fig. 3 Osculigera cf. vautrinioides Vogel, probably from Qarn Murrah, BM LL22460, Iraq Petroleum Company Collection; polished section

of right valve; x 1.

Figs 4,5 Lapeirousia sp.; 4, Jebel bu Milh, section 2, Qahlah Formation, bed 5, the acteonellid gravels; BM LL41955, conjoined pair of
right valves; 4a, ventral view; 4b, view of commissural surface; x 1; 5, Jebel Thanais, section 4, loose from lower part of Simsima
Formation; BM LL41956, conjoined pair of individuals; view showing commissural surface with pseudo-pillars below internal mould of left

valve; X 1.

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298

with thin-walled polygonal cells resembling those of Durania
rather than Biradiolites. LL41922 includes three specimens
with the upper valve in place, which is slightly concave and
smooth in the centre but develops low rounded plicae
towards the margins. LL41924 includes one specimen with
the upper, left valve present but, in this case it is hemispheri-
cally domed above the body-cavity of the right valve. We
consider this to be a phenotypic variation, possibly related to
the angle of the growth surface to the long axis of the right
valve. Right valve steeply conical, ‘D’-shaped in section, with
wide and smooth, somewhat indented radial bands on the
straight part of the ‘D’. Radial bands separated by a large
plication with two sub-plications at its crest. Rounded part of
‘D’ with approximately fourteen evenly distributed plicate
ribs. Growing surface undulates with the plication of the
outer shell sculpture.

Durania cf. apula (Parona, 1900) Pl. 6, fig. 6

1900 Biradiolites apulus Parona: 21, pl. 3, figs 1-3.Q:
1909 Sauvagesia apulus Parona; Toucas: 97, fig. 65.

MATERIAL. From Jebel Rawdah, section 2, bed 10, BM
LL41951.

DESCRIPTION. Medium sized species with approximately 40
narrow vertical ribs on lower valve. Wide growth surface of
outer shell layer of right valve, which is not plicated. Ventral
radial band relatively narrow and somewhat concave, with fine
radial striations. Dorsal radial band narrow with fine riblets.
Radial bands separated by a convex interband with a few

N.J. MORRIS AND P.W. SKELTON

sub-plicae. Growth surface of right valve has single narrow
round holes approximately 2mm in diameter at the mid-point of
its width near the position of the radial bands. Similar features
have been found in Santonian hippuritids and are being
described by Skelton & Vicens (in prep.), who regard them as
the crypts of parasitic or commensal organisms.

Pl. 6, fig. 1

MATERIAL. From Jebel Huwayyah, section 1, Qahlah gravel,
BM LL41948; two specimens from Qarn Mulayh, BM
LL41949-50, Skelton Collection.

DESCRIPTION. Large form with approximately 70 ribs on the
external surface of the right valve matched by undulations on
the wide growth surface, and prominent down-twisted con-
cave smooth dorsal radial band.

Durania form A

Fig. 6

MATERIAL. From Jebel Rawdah, section 1, bed 4, BM
LL41952; section 2, bed 6, BM LL41953; three specimens,
section 2, loose, BM LL41954.

Durania form B

DESCRIPTION. Small species with approximately 40 evenly
spaced bifurcating ribs. Outer shell layer of right valve
relatively thin with a folded growth surface, each upward fold
in the position of the interspace between the ribs.

REMARKS. The outer shell surface of the right valve and the
growth form are similar to Durania cornupastoris as figured
by Cobban, Skelton & Kennedy (1991: pls 1-3).

Fig. 6 Durania form B; a group of three conjoined individuals with the remains of a fourth on the under surface; BM LL41954, x 0-75.

PLATE 10
Fig. 1
commissural surface of right valve; x 1.

Pseudosabinia aff. klinghardti (Boehm), Jebel bu Milh, section 2, bed 7-8, ? uppermost bed of Qahlah sands; BM LL41964, view of

Figs 24 Colveraia aff. variabilis Klinghardt; 2, Qarn Mulayh, LL41958, Skelton Collection, view looking down on rim of right valve
showing the internal structure of the left valve; x 1; 3, Jebel Thanais, loose from basal beds of Simsima Formation, BM LL41961, dorsal
view of the two valves, x 1; 4, central Jebel Faiyah, loose from lower part of Simsima Formation, BM LL41960, Skelton Collection; 4a,
view looking down on internal structure of left valve; 4b, ventral view of right valve; x 1.

Fig. 5 Osculigera cf. vautriniodes Vogel, Qarn Murrah, BM LL41770, eroded section of right valve; x 1.

Fig. 6 Lapeirousia sp., Jebel Buhays, loose from basal beds of Simsima Formation, BM LL41957; minute pseudo-pillars just visible; in the
lower part of the figure the emplacement of the last shell layer is divided below the pseudopillars; x 2.

299

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS

300

Fig. 7 Lapeirousia jouanneti Des Moulins, from the Upper Campanian, near Cognac, France, BM LL41976; a, b, upper and ventral view of
a flat-based individual with pseudopillars marked, x 0-4; c, close-up of ventral pseudopillar showing its spout-like shape and division of the

last layer of outer shell around it, x 1-5.

Durania spp.

Fragmentary or poorly preserved specimens that we can
attribute to the genus Durania occur at most levels in the
Qahlah and Simsima Formations: at Jebel Huwayyah, section
1, from bed 13, the basal Simsima conglomerate, the top
rudist bed (two silicified specimens), and from near the top of
the Simsima Formation approximately 6 m above the top
oyster bed; at Jebel Rawdah, section 1, bed 2, and section 2,
bed 16; and at Qarn Mulayh, a large thick-celled form with
strangely shaped cells.

PLATE 11

Fig 1-5 Pseudosabinia aff. klinghardti (Boehm); 1, Jebel Fayah, basal Durania-facies of the Simsima Formation, BM LL41971, Skelton
Collection; eroded valve showing striate appearance of the outer part of the inner shell layer; x 0-4; 2, Virovi (between Lanac and Vestari,
near Rtanj mine), eastern Serbia; BM LL41652, paratype of Sabinia rtanjica Pejovic; section through right valvewith aréte cardinale and
ligament (arrowed); x 1-5; 3, Qarn Murrah, BM LL41972, Skelton Collection; close up of cellular structure of outer shell layer of right
valve; x 5; 4, 5, Jebel bu Milh, section 2, Qahlah Formation; 4, BM LL41963, eroded commisural surface of a right valve; x 1; 5, BM
LL41962, an eroded left valve; 5a, oblique dorsal view; 5b, posterior view; x 0-6.

N.J. MORRIS AND P.W. SKELTON

Subfamily LAPEIROUSIINAE Kuhn, 1932
Genus LAPEIROUSIA Bayle, 1878

TYPE SPECIES. Sphaerulites jouanneti Des Moulins, 1826,
Upper Campanian or Lower Maastrichtian, Aquitaine,
France.

Lapeirousia sp. PI. 9, figs 4, 5; Pl 10; fig.
MATERIAL. Two small joined specimens from Jebel Bu Milh,
section 2, from the acteonellid bed towards the top of the
Qahlah Formation, BM LL41955; three specimens, two of
which are joined, from Jebel Thanais, from scree below basal
part of the Simsima Formation, BM LL41956; one from Jebel
Buhays, section 1, LL41957.

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS

302

DESCRIPTION. Small ovoid conical species, with the right
valve approximately 75mm high and its marginal surface an
ovoid approximately 37x45 mm. The structure of the pseudo-
pillars of Lapeirousia jouanneti is shown in Fig. 7.

Genus OSCULIGERA Kuhn, 1932

TYPE SPECIES. Osculigera cleggi Kthn, 1932, by original
designation.

SYNONYM. ?=Vautrinia Milovanovié 1938 (type species,
Lapeirousia syriaca Vautrin, 1933, by original designation).

REMARKS. Species differ in degree of complication of undu-
lation of growth surface of outer shell layer.

Osculigera cf. vautrinioides Vogel, 1970 Pip shige. 3:
PIP 10; figs
cf. 1970 Osculigera vautrinioides Vogel: 69, pl. 7, figs 3, 4,

6; pl. 8, fig. 3.

MATERIAL. 21 fragmentary specimens from Qarn Murrah,
BM LL41770-80; BM LL41781—90, Skelton Collection; also
from the same locality BM LL41791 (a block with six
conjoined juveniles), and BM LL41792 (one specimen and
some fragments), Skelton Collection; also BM LL22460-63
(identified by A. Grubi¢ as O. magna Kiihn) labelled Trucial
Coast, but almost certainly the same locality, Iraq Petroleum
Co. Collection (RN 39/2,4,5.).

DESCRIPTION. Right valve varies from a low inverse cone to a
subcylinder. Surface with fine, low, even vertical ribs.
Pseudopillars project into body-cavity in low but obvious
gentle curves. Outer shell layer of right valve very thick,
about equal to the diameter of the body-cavity. Shell margin
approximately planar. Cells sub-polygonal and rather short,
inner shell layer very thin. Approximately 24 radiating undu-
lations, each topped by a row of radially elongate, radiating
projections (secondary pseudopillars of Kthn) on the
secreted marginal surface, which occasionally branch as they
approach the outer shell surface. Up to about 12 projections,
regularly spaced within each line, which are not visible in very
small individuals.

COMMENT. It is very difficult to see consistent differences
between the species described by Kuhn and Vogel. The shell
thickness and form of the radiating tuberculate undulations
seem to be very similar to the specimen figured by Vogel as
O. vautrinioides.

Subfamily JOUFIINAE Karacabey-Oztemiir 1981

GENERA INCLUDED. Colveraia (?=Dechaseauxia) and others
from Serbia and Romania, Joufia, Pseudosabinia gen. nov.
and an undescribed genus for Radiolites albonensis Toucas,
1908.

REMARKS. These are highly specialised radiolitids with pal-
lial canals and well-developed aréte cardinale. They differ
from the the subfamily Chiapasellinae Alencaster, which lack
the ligamentary invagination. The aréte cardinale of one
species of the family, Pseudosabinia rtanjica, has the split
inner ligament preserved on its inner surface (PI. 11, fig. 2).
They are fixed by the right valve.

N.J. MORRIS AND P.W. SKELTON

Genus COLVERAIA Klinghardt, 1921

TYPE SPECIES. Colveraia variabilis Klinghardt, 1921, by origi-
nal designation.

SYNONYMS. Branislavia Sladi¢-Trifunovicé, 1983a; ?Dech-

aseauxia Tavani, 1949.

Colveraia aff. variabilis Klinghardt, 1921
Pl. 10, figs 2-4

MATERIAL. One specimen from Qarn Mulayh, BM
LL41958; one specimen from Qarn Murrah, BM LL41959;
one specimen from Jebel Faiyah, north-west face, low in the
Simsima Formation, low in the Dictyoptychus-facies, BM
LL41960, all Skelton Collection; one specimen from Jebel
Thanais, loose from basal Simsima Formation, BM LL41961.

DESCRIPTION. Moderately thick calcitic outer shell layer of
the right valve with a narrow elongate aréte cardinale. Canals
obvious in the inner shell layer of both valves. In the right
valve a single row of large, subquadrate canals separate the
sockets and myophores from the outer shell layer. In the left
valve the canals are somewhat narrower and radially elongate
and penetrate the teeth and myophores. In one specimen (PI.
10, fig. 4a) the outer surfaces of both the teeth and the
myophores are longitudinally ridged, with the ridges appar-
ently formed by the canal walls, which interdigitate with
ridges on the muscle attachment surface of the opposing
valve.

Genus PSEUDOSABINIA Morris & Skelton gen. nov.
TYPE SPECIES. Pseudosabinia klinghardti (Boehm, 1927).

INCLUDED SPECIES. Sabinia klinghardti Boehm, 1927, Sab-
inia serbica Kihn & Pejovi¢, 1959, and ?Sabinia rtanjica
Pejovic, 1967.

DIAGNOSIS. Both valves extended, conical, gently curved.
Outer shell layer finely cellulo-prismatic to compact in the
right valve, thinner and smooth in the left valve. Well-
developed aréte cardinale projecting between two close,
well-developed teeth, flanked by radiolitiform myophores in
left valve. Inner shell canaliculate throughout in both valves.

REMARKS. Originally assigned to Sabinia, the species of this
genus are recognized by the finely polygonal cellulo-prismatic
structure of the outer calcitic layer of the right valve, and for
their radiolitiform myocardinal apparatus. In P. serbica the
cells appear to be present in some areas of the right valve
outer layer (Kiihn & Pejovic , 1959: figs 7, 8), whereas they
occur throughout the outer shell layer of the right valve of S.
klinghardti. The thickness and surface sculpture of coarse,
zig-zag growth rugae of this outer shell layer are similar in S.
serbica and §. klinghardti.

In the interpretation of Philip (1986) all species of Sabinia
are considered to be related and are off-shoots of the
Radiolitidae that have secondarily reduced the outer calcitic
shell layer of the right valve. We consider that there may be
at least two distinct genera within the genus Sabinia as
presently constituted, and that these may be quite unrelated.
The features of the true Sabinia with its type species, Sabinia
anienis Parona 1908, may relate it to the Plagioptychidae.

NOTE ON THE TYPE SPECIES OF SABINIA. Parona described
three species: in sequence they were S. sublacensis, S. sinuata

LATE CAMPANIAN-MAASTRICHTIAN RUDISTS

and S. anienis from the Pietra di Subiaco, Late Cretaceous, of
Monte Affilano, Valle dell’Aniene, Provincia di Roma (the
type specimens are in Rome). All three nominal species may
be merely varieties. In Parona’s material a septum cuts off the
posterior accessory cavity from the body-cavity in the left
valve (Parona, 1908: figs a, c). It runs between the anterior
tooth and the postero-ventral margin, and could be equiva-
lent to the septum in the left valve of the plagioptychids.
Parona does not describe the structure of the outer shell layer
of the right valve in any detail.

Pseudosabinia aff. klinghardti (Boehm, 1927) Pl. 10,
fig. 1; Pl. 11, figs 1-5

Sabinia klinghardti Boehm: 205, pl. 15, figs 1, 2;
pl. 16, fig. 1.

aff. 1927

?aff. 1927 Schiosia bilinguis Boehm: 207, pl. 18, figs la—Ic.

aff. 1967 Pseudosabinia rtanjica Pejovic : 295-97, pl. 1,
fig. 1.

aff. 1986 Sabinia rtanjica tunisiensis Philip: 248, 49, pl. 1,
figs 1-6.

Types. The holotype, BM L49455 from the Campanian-
Maastrichtian, east of Hereke and west of Tauchanly,
Bythinia, north-west Turkey, has both valves preserved; the
paratype, BM 49454 from the same locality, is a much larger
specimen of a crushed right valve. The holotype of Schiosia
bilinguis, BM L50929, is from the ‘Upper Senonian’ between
Herake and Tauchanly, Bithynia. BM LL41652 is an unfig-
ured paratype of P. rtanjica Pejovic, from near Rtanj mine,
eastern Serbia.

NEW MATERIAL. From Qarn Murrah, a large crushed right
valve with most of the outer shell layer missing except for a
thin layer remaining, together with a second crushed valve
which may be left or right, BM LL41972, Skelton Collection;
four specimens from Jebel Bu Milh, section 2, Qahlah Sands,
BM LL41962-65; two possible specimens from Jebel Raw-
dah, section 4, bed 1, the basal rudist bed, BM LL41966-67,
and another small left valve that shows well-developed pallial
canals and a narrow aréte cardinale, BM LL41968; a large left
valve from the north-west face of Jebel Faiyah, Simsima
Formation, the Durania-facies, BM LL41971, and a small left
valve from central Jebel Faiyah, BM LL41981, both Skelton
Collection.

DESCRIPTION. The material from Qarn Murrah is badly
crushed but does show the cellular outer shell layer (PI. 11,
fig. 3). The large, horn-like right valve (PI. 11, fig. 4), at least
16 cm long, has a long narrow aréte cardinale, a thick inner
shell consisting of irregularly polygonal pallial canals, fine at
the valve margin, becoming coarser inwards, and prominent
cystose tabulae. Accessory cavities are apparently absent.
There are two sockets for prominent prong-like teeth with
longitudinal grooves and ridges either side of a low, ridge-like
tooth, immediately ventral to the aréte cardinale. The myo-
cardinal arrangement is not capriniform. The small right
valve of Pl. 10, fig. 1 shows the aréte cardinale straddled by
the ridged sockets for the teeth. Two equally prominent teeth
are present in the fixed left valve of Pl. 11, fig, 5.

REMARKS. The progression from small to larger pallial canals
in the left valve of S. bilinguis Boehm is similar to the pattern
in the present species, otherwise Boehm’s species has very
few characters preserved. The outer shell layer of the right
valve of Pseudosabina rjtanjica is thin (less than 1 mm, fide

303

Philip) and of dense fibrous calcite, ie not cellular as in P.
klinghardti. This seems to be the only difference, but is
confirmed by our material. Pejovi¢ (1967) and Philip (1986)
suggest that a row of canals separating the myophores from
the body-cavity distinguishes P. rtanjica from P. klinghardti.
Careful inspection of Boehm’s holotype leads us to the
conclusion that this difference does not exist. The myophores
actually have a purely radiolitiform arrangement, albeit with
contained canals, and face outwards opposing the inner
surface of the right valve.

Pejovic’s (1967) description of Sabinia rtanjica was based
only on eight specimens of the upper, left valve, in which the
myophores were stated to be separated from the living
chamber by a number of pseudocanals, characterized by
regularly spaced tabulae and not considered homologous with
the canals of the Caprinidae. A well- preserved small left
valve collected by one of us (PWS) from the ?Lower Campa-
nian of Monte Kamilja, near Leposavi¢ in south-west Serbia,
confirms the description of P. rtanjica in having large canals
on inner sides of the myophores. However, this feature is in
fact the coarse canaliculate structure of the myophoric apo-
physes and is also present in the holotype of Pseudosabinia
klinghardti. Philip’s (1986) Tunisian form differs from Pejov-
ic’s original description in the length of the aréte cardinale
which reaches well between the teeth, unlike that in the
Serbian original which, it is claimed, only reaches the top of
the teeth/sockets. Inspection of Pejovié ‘s 1967, pl. 1, fig. 1
suggests that this is incorrect. Secondly Philip stated that the
pallial canals of the lower (right) valve are of oval section in
the Tunisian form, but oval and polygonal in the nominal
subspecies. He also claimed that the upper valve canals have
a suboval section in the Tunisian form, but these are pentago-
nal, hexagonal or triangular in the nominal subspecies. We
suspect that these latter two distinctions reflect only the
diagenetic growth of fibrous cement crust within the canals of
the Tunisian material.

Pseudosabinia rtanjica has a very thin, compact outer shell
layer in the right valve, according to Philip (1986), which is in
contrast to the much thicker outer shell layer of Pseudosab-
inia klinghardti, although there are dense areas in the outer
part of this layer in Boehm’s holotype. This seems to be
supported by our rather poorly preserved material, although
a specimen with part of both valves preserved, apparently of
P. rtanjica, from Serbia, kindly donated from Mdm. Pejovic,
shows a thin zone of cells on the innermost zone of the outer
shell layer grading into dense structure in the outer part. This
suggests the possibility of intergradation between the two
taxa. Inspection of further material would be required to
confirm this.

ACKNOWLEDGEMENTS. We would like to thank Diana Clements for
considerable help in both sorting the material and preparing the
manuscript. We are indebted to Phil Crabb for taking the photo-
graphs, and to Tony Wighton and his colleagues for cutting the
sections of the rudists.

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305

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Note added in proof: One of us (PWS) has recently had the
Opportunity to see some specimens of Durania mutabilis
(Stoliczka, 1871) collected from the Maastrichtian of south-
ern India by Professor Malcolm Hart. It is clear that the
material described above (p. 296) as Durania cf. gaensis
(Dacqué, 1903) can be attributed to D. mutabilis. Whether
D. gaensis and D. mutabilis (which has priority) should be
considered to be synonyms remains to be resolved.

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