Acta Geologica Polonica, Vol. 62 (2012), No. 1, pp.

6389

Late CenomanianTuronian (Cretaceous) ammonites from

Wadi Qena, central Eastern Desert, Egypt: taxonomy, biostratigraphy and palaeobiogeographic implications
EMAD NAGM1 AND MARKUS WILMSEN2
Geology Department, Faculty of Science, Al-Azhar University, Assiut, Egypt.
E-mail: emad.nagm@yahoo.com
2
Senckenberg Naturhistorische Sammlungen Dresden, Museum fr Mineralogie und Geologie,
Sektion Palozoologie, Knigsbrcker Landstr. 159, D-01109 Dresden, Germany. `
E-mail: markus.wilmsen@senckenberg.de
1

ABSTRACT:
Nagm, E. and Wilmsen, M. 2012. Late CenomanianTuronian (Cretaceous) ammonites from Wadi Qena, central
Eastern Desert, Egypt: taxonomy, biostratigraphy and palaeobiogeographic implications. Acta Geologica Polonica,
62 (1), 6389. Warszawa.
In Egypt, marine Upper CenomanianTuronian strata are well exposed in the Eastern Desert. The southernmost
outcrops are located in the central part of Wadi Qena, where the lower Upper Cretaceous is represented by the fossiliferous Galala and Umm Omeiyid formations. From these strata, numerous ammonites have been collected
bed-by-bed and 13 taxa have been identified, which are systematically described herein. Four of them (Euomphaloceras costatum, Vascoceras globosum globosum, Thomasites gongilensis and Pseudotissotia nigeriensis) are
recorded from Egypt for the first time. The ammonite ranges are used for a biostratigraphic zonation of the lower
Upper Cretaceous succession in the northern and central part of Wadi Qena: the Upper CenomanianLower Turonian has been subdivided into five biozones (including a new upper Lower Turonian biozone based on the occurrence of Pseudotissotia nigeriensis), and one biozone has been recognized in the Upper Turonian.
Palaeobiogeographically, the ammonite assemblage has a Tethyan character. During the Early Turonian, influences
of the Vascoceratid Province were predominant with strong affinities to typical Nigerian faunas. This shows the
significance of faunal exchange between Egypt and Central and West Africa via the Trans-Saharan Seaway. Compared to contemporaneous ammonoid faunas from the northern part of the Eastern Desert, Boreal influences are
much less obvious in Wadi Qena. Thus, the present study greatly enhances the knowledge of the Late Cretaceous
palaeobiogeography and biostratigraphy of Egypt and adjacent areas.
Key words: Upper Cretaceous; Ammonites; Systematic palaeontology; Biostratigraphy;

Palaeobiogeography; Egypt.

INTRODUCTION
Marine lower Upper Cretaceous successions are
well exposed in the northeastern part of Egypt (Sinai
and Eastern Desert; Text-fig. 1). The southernmost
exposures of Upper CenomanianTuronian sediments

are preserved in the central part of the Eastern Desert

in Wadi Qena, and ammonites are among the most important faunal elements in these strata. However, only
three papers from the last century have dealt with this
topic (Eck 1914; Douvill 1928; Luger and Grschke
1989). Therefore, this paper aims to enhance the

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EMAD NAGM AND MARKUS WILMSEN

knowledge of the taxonomy of the Late CenomanianTuronian ammonite fauna in Wadi Qena by means
of a modern systematic approach and description of a
large fauna of ammonites collected bed-by-bed from
the Galala and Umm Omeiyid formations. In total, 90
ammonites have been collected during two field cam-

paigns (April and June 2011) from two sections in

Wadi Qena, forming the basis of the study. In addition,
a detailed biostratigraphic framework is proposed for
the succession based on the ranges of the ammonites.
Finally, the palaeobiogeographic affinities of the ammonite assemblage are briefly discussed.

Text-fig. 1. Geographical and geological framework. A: map of Egypt with position of Wadi Qena (B); B map of Wadi Qena with position of study area (C); C: geological
map of study area (modified after Conoco, 1987)

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CENOMANIAN TURONIAN AMMONITES FROM EGyPT

GEOLOGICAL SETTING
Wadi Qena is located in the central part of the
Egyptian Eastern Desert (Text-fig. 1A, B). The main
part of Wadi Qena is about 200 km long and runs from
the southern slopes of the Southern Galala Plateau,
near the Gulf of Suez, towards the south southeast. Near
the town of Qena City it grades into the River Nile valley. Wadi Qena is bordered by the uniform, elevated and
flat-topped Eocene limestone Plateau in the west and
the serrated, varicoloured basement rocks of the Red
Sea Mountains range in the east (Text-fig. 1B). Wadi
Qena clearly developed along faults that run parallel to
the Gulf of Suez and the Red Sea Mountains, which are
well exposed at numerous places within the wadi, especially on the eastern side (Bandel et al. 1987). Sedimentary successions are well exposed on the western
flank of the wadi while Quaternary alluvial fans predominate in the centre (Text-fig. 1C).
The sedimentary rocks of Wadi Qena range in age
from Palaeozoic to Quaternary (Text-fig. 1C). Most of
the Palaeozoic rocks are non-marine and well developed in the northern part of the wadi. The distribution
of the Palaeozoic sediments was more or less controlled by older, pre-Cenomanian structural events
(Said 1990; Issawi et al. 1999). The first marine transgression took place during the Late Cenomanian. This
resulted in the deposition of shallow marine, mixed siliciclastic and carbonate sediments. During Late CenomanianTuronian times, the southern shoreline of the
Tethyan seaway was situated immediately south of the
study area. The marine transgression continued with
some phases of regression during the later parts of the
Late Cretaceous and Tertiary, leaving Coniacian
Eocene strata in the study area (see Text-fig. 1C).
SECTIONS
Two sections, located on the western margin in the
northern part of Wadi Qena (Text-fig. 1C), have been
measured bed-by-bed. One section, measured at N 27
57` 57``, E 32 30` 58``, is termed the north Wadi Qena
section (Text-fig. 2). The other section, located 25 km
south of the north Wadi Qena section, was measured at
the mouth of Wadi El-Burga entering Wadi Qena at N
27 44` 51``, E 32 33` 55`` and is consequently termed
the Wadi El-Burga section (Text-fig. 2).
The sedimentary succession starts with the Palaeozoic Naqus Formation (non-marine, kaolinitic,
medium- to coarse-grained sandstones; see Abdallah et
al. 1992 and Issawi et al. 1999) which, in places,
overlies basement rocks unconformably. Along a trun-

cated palaeosol, the Naqus Formation is overlain by

the Galala Formation. The Galala Formation (Text-fig.
2) starts with silty, glauconitic sandstones rich in the
trace fossil Thalassinoides isp. and fragmented mollusc shells, indicating shallow marine conditions.
Higher up-section, shale beds are common (especially
in the north), with a few sandy limestone and thin bioturbated sandstone intercalations. The lower part of the
Galala Formation (35 m in the north and 25 m in the
south) is terminated by a fossiliferous limestone bed,
yielding a Late Cenomanian faunal assemblage including Inoceramus ex gr. pictus Sowerby (Nagm et
al. 2011). The following interval consists of gypsiferous shale in the north and silty shale with thin sandstone interbeds in the south. It contains a highly fossiliferous limestone bed around the ~55-m-level (from
the base of the Galala Formation) in both sections.
This bed is characterized by a typical Late Cenomanian fauna represented by oysters [Ceratostreon
flabellatum (Goldfuss), Curvostrea rouvillei (Coquand), Ilymatogyra africana (Lamarck)], nautiloids
[Angulithes mermeti (Coquand)], ammonites [Neolobites vibrayeanus (dOrbigny)] and echinoids (Hemiaster cubicus Desor) indicating a fully marine, lagoonal environment (cf. Nagm 2009; Wilmsen and
Nagm 2012). Marls with calcareous nodules are common in the following part, which ends at a sharp and
more or less irregular surface at 70 m and 64 m respectively in the two sections. The upper part of the
Galala Formation (50 m in the north and 45 m in the
south) is characterized by a high carbonate content
and consists of thick limestone beds intercalating with
marl containing calcareous nodules and/or sandy siltstone (in the south). These sediments are highly fossiliferous, containing abundant latest Cenomanian and
Early Turonian ammonites (most of the studied specimens have been collected from this interval). The
Galala Formation is truncated along a prominent unconformity at the base of the Umm Omeiyid Formation. This unconformity has also been recognized in
Wadi Araba and placed in the LowerMiddle Turonian boundary interval (Nagm et al. 2010a, b). The
thickness of the Galala Formation is about 120 m in
the North Wadi Qena section and about 112 m in the
Wadi El-Burga section.
The Umm Omeiyid Formation overlies the Galala
Formation unconformably, attaining a thickness of
about 50 m in both sections. It consists of unfossiliferous fissile siltstone with thin, medium-grained, in
part hummocky cross-stratified sandstone intercalations. The upper part of the formation is also characterized by siliciclastic facies, but additionally contains
a few fossiliferous limestone beds and/or marl inter-

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EMAD NAGM AND MARKUS WILMSEN

Text-fig. 2. Graphic logs of the North Wadi Qena and Wadi El-Burga sections with stratigraphic distribution of the zonal ammonite markers. The dashed lines represent
inferred ranges of taxa depending on other stratigraphic data. For location see Text-fig. 1C

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CENOMANIAN TURONIAN AMMONITES FROM EGyPT

calations. The calcareous layers yield abundant early

Late Turonian ammonites [Coilopoceras requienianum (dOrbigny)]. The Umm Omeiyid Formation is
overlain by the Coniacian Taref Sandstone Formation
along a sharp, ferruginous surface.
SySTEMATIC PALAEONTOLOGy
Repositories of specimens: 90 specimens (mostly
composite internal moulds) form the basis of the study.
Illustrated material is kept in the Museum fr Mineralogie und Geologie (MMG) of the Senckenberg
Naturhistorische Sammlungen Dresden (SNSD), Germany (repository AfK). Additional specimens are
housed in the Department of Geology, Al-Azhar University, Egypt (repository AUEN).
Dimensions: All linear dimensions, taken with a
Vernier Caliper, are given in millimetres. The abbreviations that are used in the present work are: maximum diameter (D), whorl breadth (Wb), whorl height
(Wh), breadth of umbilicus (U). Figures in parentheses are dimensions as a percentage of diameters.
Suture terminology: The suture terminology follows Wedekind (1916) as applied by Kullman and
Wiedmann (1970) and revised in parts by Korn et al.
(2003): E=external lobe, A=adventive lobe (=lateral
lobe, L, of former nomenclature), U=umbilical lobe,
I=internal lobe.
Systematics: The ammonites studied are arranged
systematically according to Wright et al. (1996). The
terminology used for the description of the taxa follows the glossary in the Treatise on Invertebrate Paleontology, Part L, Mollusca 4 (Wright et al., 1996). In
order to keep this article as short as possible, the synonymy lists only contain a few selected references
which are important for the discussion.

Order Ammonoidea von Zittel, 1884

Suborder Ammonitina Hyatt, 1889
Superfamily Hoplitoidea Douvill, 1890
? Family Engonoceratidae Hyatt, 1900
Genus Neolobites Fischer, 1882
TyPE SPECIES: Ammonites vibrayeanus dOrbigny,
1841; by original designation.
Neolobites vibrayeanus (dOrbigny, 1841)
(Text-figs 3, 6A)
1841. Ammonites vibrayeanus dOrbigny, 322, pl. 96, figs
13.
1981. Neolobites vibrayeanus (dOrbigny); Kennedy and
Juignet, 23, figs 34, 6a; text-fig. 5.
1989. Neolobites vibrayeanus (dOrbigny); Luger and
Grschke, 366, pl. 39, fig. 3, text-fig. 5.
2005. Neolobites vibrayeanus (dOrbigny); Wiese and
Schulze, 933, figs 49. [with synonymy]
2010a. Neolobites vibrayeanus (dOrbigny); Nagm et al.,
475, figs 4, 5AF.

MATERIAL: Twenty specimens (AfK 297301, AUEN

115) from three different levels in the middle part of the
Galala Formation in both sections (Text-fig. 2).
MEASUREMENTS:
Specimen
AfK 298
AfK 299
AfK 300
AfK 301

D
86.4 (100)
92.9 (100)
57.3 (100)
72.5 (100)

Wb
20.6 (23.8)
20.2 (21.7)
16.1 (28.0)
18.3 (25.2)

Wh Wb/Wh U
51.0 (59.0) 0.40 8.1 (9.3)
51.7 (55.6) 0.39 8.8 (9.4)
33.7 (58.8) 0.47 4.9 (8.5)
46.9 (64.6) 0.39 4.4 (6.0)

DESCRIPTION: Most of the specimens are fully septate internal moulds with very involute coiling. The

Text-fig. 3. Whorl section and suture line of Neolobites vibrayeanus (dOrbigny, 1841), Upper Cenomanian of the Galala Formation, specimen AfK 301

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EMAD NAGM AND MARKUS WILMSEN

small umbilicus, comprising about 6% of the diameter,

is shallow, with low and rounded umbilical walls. The
flanks are slightly convex with a narrow and flat venter. The whorl section (Text-fig. 3) is highly compressed, elevated, and has a lanceolate shape with
greatest breadth around mid-flank. The suture lines
(Text-fig. 3) are poorly preserved and characterized by
broad saddles with somewhat narrower and lanceolate
lobes that become shallower towards the umbilicus.
The ornamental features are also poorly preserved except for some delicate clavi at the ventrolateral margin
(e.g., Text-fig. 6A45).
REMARKS: Despite the imperfect preservation, the
specimens collected from the Galala Formation of
Wadi Qena can be assigned to N. vibrayeanus without any doubt. Neolobites includes several species
and many morphological variants, the differences between which have been discussed by Kennedy and
Juignet (1981) and Wiese and Schulze (2005). Most,
if not all, Neolobites specimens that have been
recorded from Egypt are representatives of one
species, i.e., N. vibrayeanus. The morphological
variation in the Egyptian material (presence or absence of very weak ribs and ventrolateral clavi) is related to different taphonomic conditions (for more
details on N. vibrayeanus from Egypt see Nagm et
al. 2010a).
OCCURRENCE: Neolobites vibrayeanus is a widespread and characteristic early Late Cenomanian
species, occurring in France, Spain, Portugal, Morocco, Algeria, Tunisia, Egypt, Israel, Lebanon, Syria,
Saudi Arabia, Oman, Niger, Bolivia, Colombia, Peru
and Venezuela. According to Wiese and Schulze
(2005), the species preferred shallow-water environments in subtropical/tropical areas. In the study area,
N. vibrayeanus characterizes the lower Upper Cenomanian Neolobites vibrayeanus Zone, which is (as
defined by Nagm et al. 2010b) equivalent to the Cenomanian standard ammonite Zone of Calycoceras
(Proeucalycoceras) guerangeri (cf. Kennedy 1984;
Wright et al. in Wright and Kennedy 1984; Hancock
1991).
Superfamily Acanthoceratoidea de Grossouvre, 1894
Family Acanthoceratidae de Grossouvre, 1894
Subfamily Euomphaloceratinae Cooper, 1978
Genus Euomphaloceras Spath, 1923
TyPE SPECIES: Ammonites euomphalus Sharpe,
1855; by monotypy.

Euomphaloceras costatum Cobban, Hook and

Kennedy, 1989
(Text-figs 4, 6B)
1989. Euomphaloceras costatum Cobban, Hook and
Kennedy, 37, figs 37, 77SEE, 78AH.
2005. Euomphaloceras costatum Cobban, Hook and
Kennedy; Gale et al.

MATERIAL: Three specimens (AfK 305307) from

the upper part of the Galala Formation (97 m above the
base) at the Wadi El-Burga section (Text-fig. 2).
MEASUREMENTS:
Specimen
D
Wb
Wh
Wb/Wh
U
AfK 306 64.8 (100) 32.3 (49.8) 25.7 (39.6) 1.25 23.8 (36.7)

DESCRIPTION: The coiling is evolute, with a large

umbilicus and rounded umbilical walls. The whorl
section (Text-fig. 4) is quadratic, with the greatest
breadth at the umbilical tubercles. Flanks are narrow,
flattened and ribbed, with strong umbilical and ventrolateral tubercles occurring on all main ribs. The
venter is broad and slightly convex, showing a weak
ridge at the site of the siphuncle, and carries three
rows of blunt tubercles (the middle one on the siphonal
ridge). The suture line is poorly preserved.

Text-fig. 4. Whorl section of Euomphaloceras costatum Cobban, Hook and

Kennedy, 1989, Upper Cenomanian of the Galala Formation, specimen AfK 305

REMARKS: The specimens described herein are very

close to the holotype as described by Cobban et al.
(1989). They differ in their less dense ribbing. In spite
of their imperfect preservation, they are assigned to E.
costatum without reservation.
OCCURRENCE: Upper Cenomanian of New Mexico
(USA), southern England and Brazil. In the study area,
the species has been collected from the upper part of
the Vascoceras cauvini Zone, which is (as defined by
Nagm et al. 2010b) equivalent to the Neocardioceras
juddii Zone, in the standard ammonite zonation
(Kennedy 1984; Wright et al. in Wright and Kennedy
1984; Hancock 1991). This is the first record of E.
costatum from Egypt, which confirms the temporal

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CENOMANIAN TURONIAN AMMONITES FROM EGyPT

equivalence of the V. cauvini Zone to the N. juddii

Zone (see below).
Genus Pseudaspidoceras Hyatt, 1903
TyPE SPECIES: Ammonites footeanus Stoliczka,
1864; by original designation.
Pseudaspidoceras sp.
(Text-figs 5, 6C)
MATERIAL: One specimen (AfK 304) from the upper
part of the Galala Formation (95 m above the base) at
the Wadi El-Burga section (Text-fig. 2).
MEASUREMENTS:
Specimen
D
Wb
Wh Wb/Wh
U
AfK 304 95.0 (100) 36.9 (38.8) 53.5 (56.3) 0.68 29.3 (30.8)

DESCRIPTION: The coiling is moderately evolute,

with a large umbilicus (ca. 30%). The whorl section
(Text-fig. 5) is trapezoidal, with the strongest inflation at the umbilical shoulders. From the umbilical
shoulders, the flanks converge towards a feebly convex venter with narrowly rounded ventrolateral
shoulders. The ornamentation consists of distant,
rounded and sometimes branched ribs carrying umbilical bullae. Each rib ends at the ventrolateral shoulder with a strong tubercle. Ribbing weakens towards
the outer whorl. The suture line (Text-fig. 5) is well
preserved and characterized by a broad adventive
lobe.

REMARKS: This well preserved specimen has all the

diagnostic characters of the genus Pseudaspidoceras.
However, it is difficult to identify at the specific level
because of the inadequate match of its morphological
characters with those of known species of the genus.
In fact, there are only two species that come close to
the present specimen, i.e., P. conciliatum Stoliczka var.
ampakabensis Collignon (1965, p. 36, fig. 1669) and
P. pseudonodosoides Choffat (1898, p. 65, pl. 16, figs
58; pl. 22, figs 3233). However, P. conciliatum
Stoliczka var. ampakabensis differs from our specimen
in a slightly concave venter and highly indented sutures; furthermore, it is known from the Lower Turonian whereas the specimen from Wadi Qena was collected from uppermost Cenomanian strata. In
stratigraphic position, P. pseudonodosoides matches
our specimen, but it differs morphologically in its depressed whorl section with a very broadly rounded
venter. The Wadi Qena specimen possibly represents
a new species of the genus Pseudaspidoceras, but this
conclusion cannot be based on a single specimen.
OCCURRENCE: The specimen was collected from
the upper Galala Formation at the Wadi El-Burga section, 2 m above the level with Vascoceras cauvini and
2 m below the level with Euomphaloceras costatum.
Therefore, its stratigraphic position is equivalent to the
upper Upper Cenomanian Neocardioceras juddii standard ammonite Zone (sensu Wright et al. in Wright and
Kennedy 1984).
Family Vascoceratidae Douvill, 1912
Genus Vascoceras Choffat, 1898

Text-fig. 5. Suture line and whorl section of Pseudaspidoceras sp., Upper Cenomanian of the Galala Formation, specimen AfK 304

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EMAD NAGM AND MARKUS WILMSEN

Text-fig. 6. A Neolobites vibrayeanus (dOrbigny, 1841) (A1, A2 - lateral and ventral views of AfK 298, 1; A3 - lateral view of AfK 300, 1; A4, A5 - lateral
and ventral views of AfK 301, 1). B Euomphaloceras costatum Cobban, Hook and Kennedy, 1989 (B1, B2 - lateral and ventral views of AfK 305, 1).
C Pseudaspidoceras sp. (C1, C2 - lateral and ventral views of AfK 304, 1). All specimens from the Upper Cenomanian of the Galala Formation

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CENOMANIAN TURONIAN AMMONITES FROM EGyPT

TyPE SPECIES: Vascoceras gamai Choffat, 1898;

by the subsequent designation of Diener, 1925.
Vascoceras cauvini Chudeau, 1909
(Text-figs 7, 9A)
1909. Vascoceras cauvini Chudeau, 67, pl. 1, figs 1a, 2a; pl.
2, figs 3, 5; pl. 3, figs 1b, 2b, 4.
1969. Paravascoceras cauvini (Chudeau); Freund and Raab,
20, pl. 3, figs 13; text-fig. 5a, b.
1989. Vascoceras cauvini Chudeau; Luger and Grschke,
374, pl. 40, figs 3, 6, 8, 9; pl. 41, figs 14; pl. 42, fig.
1; text-figs 6g,h, 8c.
2010a. Vascoceras cauvini Chudeau; Nagm et al., 483, figs
9AB, 10B.

MATERIAL: Two specimens (AfK 302, 303) from the

upper part of the Galala Formation (93 m above the
base) at the Wadi El-Burga section (Text-fig. 2).
MEASUREMENTS:
Specimen
D
Wb
Wh Wb/Wh
U
AfK 302 96.3 (100) 41.3 (42.8) 36.1 (37.4) 1.14 33.4 (34.6)
AfK 303 98.0 (100) 34.3 (35.0) 41.5 (42.3) 0.82 37.1 (37.8)

DESCRIPTION: Both specimens are fragmentary internal moulds with relatively evolute coiling as well as
a deep and wide umbilicus (ca. 35%). The umbilical
shoulders are feebly rounded with vertical walls.
Flanks are short and convex with a broadly rounded to
nearly flat venter (in the latter case, faintly defined ventrolateral shoulders are present). The whorl section is
compressed to slightly depressed (Text-fig. 7, specimen AfK 302). The ornamentation is not preserved in
specimen AfK 302 (Text-fig. 9A34), while very
weak, shallow ribs, especially towards the venter, occur in specimen AfK 303 (Text-fig. 9A12, a fragment
of a body chamber). The suture line, characterized by
a deep external lobe and a wide, deep adventive lobe,
is clearly visible in specimen AfK 302 (see Text-fig. 7).

REMARKS: In Vascoceras cauvini two forms are

known (see Chudeau 1909); i.e. the smooth form
(e.g. Luger and Grschke 1989; Zaborski 1996; Aly
et al. 2008; Barroso-Barcenilla and Goy 2010; Nagm
et al. 2010a); and the ribbed form (e.g., Freund and
Raab 1969; Zaborski 1990, 1996; Meister et al.
1992). Two different forms also occur in the study
area: specimen AfK 302 is a typical representative of
the smooth form (Text-fig. 9A34) while specimen
AfK 303 (Text-fig. 9A12) shares characters of both
the smooth and ribbed forms in terms of dimensions
and suture line. On the other hand, it has its own characters, i.e., a flat venter and the absence of the pronounced ornamentation typical of the ribbed form
(very weak, shallow ribs occur only at the venter).
Therefore, this specimen is considered here as a transitional morph between the smooth and ribbed form
of V. cauvini.
Vascoceras pioti (Peron and Fourtau, 1904; in
Fourtau, 1904) likewise has a flat venter, but its
umbilicus is very small and the suture is different:
the lobes of V. cauvini are deeper and wider than
those of V. pioti, especially the A-lobe (Text-fig. 7).
In addition, the suture line of V. cauvini has three
saddles, while that of V. pioti has four (see Freund
and Raab 1969, p. 30, text-fig. 6f). The sutures of V.
cauvini from Wadi Qena (Text-fig. 7) are identical
to sutures in the type material as illustrated by
Chudeau (1909). Furthermore, V. pioti is from a
higher stratigraphic level, coming from the Lower
Turonian.
OCCURRENCE: Vascoceras cauvini is widely known
from the Upper Cenomanian of Nigeria, Niger, Angola,
Morocco, Algeria, Egypt, Israel, Spain and Peru. In the
study area, Vascoceras cauvini characterizes the uppermost Cenomanian Vascoceras cauvini Zone, which
is (cf. Nagm et al. 2010b) equivalent to the Neocardioceras juddii Zone in the standard ammonite zonation of Kennedy (1984), Wright et al. in Wright and
Kennedy (1984) and Hancock (1991).

Text-fig. 7. Whorl section and suture line of Vascoceras cauvini Chudeau, 1905, Upper Cenomanian of the Galala Formation, specimen AfK 302

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EMAD NAGM AND MARKUS WILMSEN

Vascoceras globosum globosum (Reyment, 1954)

(Text-figs 8, 9B)
1954. Pachyvascoceras globosum Reyment, 259, pl. 3, fig.
3; pl. 5, fig. 4; text-figs 3e, 7.
1957. Vascoceras globosum globosum (Reyment); Barber,
21, pl. 7, figs 1a, b; pl. 9, figs 4a, b; pl. 28, figs 1, 2.
2005. Vascoceras globosum globosum (Reyment); Gale et
al., 182, fig. 9.

MATERIAL: A single specimen (AfK 308) from the

upper part of the Galala Formation (102 m above the
base) at the Wadi El-Burga section (Text-fig. 2).
MEASUREMENTS:
Specimen
D
Wb
Wh Wb/Wh
U
AfK 308 80.3 (100) 69.2 (86.1) 41.5 (51.6) 1.66 10.1 (12.5)

DESCRIPTION: The specimen is a completely smooth,

globose, very involute phragmocone with a maximum
preserved diameter of 80.3 mm. The umbilicus is small
(12.5% of diameter) and very deep, with a narrowly
rounded umbilical shoulder. The flanks are short and
strongly convex, grading without ventrolateral shoulders into a broadly rounded venter. The whorl section
(Text-fig. 8) is depressed and globose, with a whorlbreadth to whorl-height ratio of 1.66. The suture line is
well preserved and illustrated in Text-fig. 8.
REMARKS: Barber (1957) erected five subspecies of
Vascoceras globosum based on variations in the degree
of inflation. Of those subspecies, V. globosum globosum
is the most inflated. The specimen from Wadi Qena is
strongly depressed and globose in shape, with a small
and deep umbilicus. These characters are diagnostic of
Vascoceras globosum globosum as described and illustrated by Barber (1957) and Zaborski (1996).
OCCURRENCE: Uppermost Cenomanian of Nigeria
and Lower Turonian of Brazil. In the study area, it has

been recorded from the Lower Turonian, i.e., the lower

part of the Choffaticeras spp. Zone. This zone is (as defined by Nagm et al. 2010b) equivalent to the upper
part of the standard ammonite Zone of Watinoceras
coloradoense (Kennedy 1984; Wright et al. in Wright
and Kennedy 1984) or Watinoceras spp. (Hancock
1991). This is the first record of Vascoceras globosum
globosum from Egypt.
Vascoceras kossmati Choffat, 1898
(Text-figs 9C, 10, 13A)
1898. Vascoceras kossmati Choffat, 63, pl. 13, figs 8, 9; pl.
14, figs 1, 2; pl. 21, figs 26, 27.
1992. Paravascoceras kossmati (Choffat); Thomel, 225, pl.
119, figs 2, 3.
2010. Vascoceras kossmati Choffat; Barroso-Barcenilla and Goy,
219, pl. 5, figs FH; text-fig. 6B. [with synonymy].

MATERIAL: Four specimens (AfK 309312) from the

upper part of the Galala Formation (102 m above the
base) at the Wadi El-Burga section (Text-fig. 2).
MEASUREMENTS:
Specimen
D
AfK 309 65.5 (100)
AfK 310 52.5 (100)
AfK 311 59.9 (100)
AfK 312 66.4 (100)

Wb
48.0 (73.2)
39.6 (75.4)
46.4 (77.4)
45.6 (68.6)

Wh Wb/Wh
U
30.2 (46.1) 1.58 13.2 (20.1)
27.4 (52.1) 1.44 9.0 (17.1)
33.0 (55.0) 1.40 7.8 (13.0
35.2 (53.0) 1.29 9.1 (13.7)

DESCRIPTION: All specimens are smooth internal

moulds characterized by strongly involute coiling. The
umbilicus is narrow and deep, showing vertical walls
and sharp margins. The flanks are strongly convex,
grading into a broadly rounded venter. Some specimens (AfK 311, 312) have wide and flat venters that
are clearly separated from the flanks by ventrolateral
shoulders (Text-fig. 13A2). The whorl section (Textfig. 10) is depressed, with an incipiently triangular

Text-fig. 8. Whorl section and suture line of Vascoceras globosum globosum (Reyment, 1954), Lower Turonian of the Galala Formation, specimen AfK 308

73
CENOMANIAN TURONIAN AMMONITES FROM EGyPT

Text-fig. 9. A Vascoceras cauvini Chudeau, 1909 (A1, A2 - ventral and lateral views of AfK 303, 1; A3, A4 - lateral and ventral views of AfK 302, 1; Upper
Cenomanian of the Galala Formation). B Vascoceras globosum globosum (Reyment, 1954) (B1, B2 - lateral and ventral views of AfK 308, 1; Lower Turonian
of the Galala Formation). C Vascoceras kossmati Choffat, 1898 (C1, C2 - lateral and apertural views of AfK 309, 1; Lower Turonian of the Galala Formation)

74

EMAD NAGM AND MARKUS WILMSEN

Text-fig. 10. Whorl section and suture line of Vascoceras kossmati Choffat, 1898, Lower Turonian of the Galala Formation, specimen AfK 309

shape. The suture line (Text-fig. 10) is simple and

characterized by a wide and deep adventive lobe.
REMARKS: Barroso-Barcenilla and Goy (2010) discussed the similarities between Vascoceras kossmati
and Vascoceras hartti (Hyatt). They regarded the subspherical shape, the small umbilical width and the depressed whorl section as the key characters of V. kossmati. The material from Wadi Qena is very close to the
Spanish material, and we thus assign it to Choffats
species without hesitation.
OCCURRENCE: Lower Turonian of Portugal, Spain
and France (see Barroso-Barcenilla and Goy 2010).
Eck (1914) noted the species from Egypt without illustration. Therefore, this is the first properly documented record from Egypt. In the study area, it occurs in the Lower Turonian, in the lower part of the
Choffaticeras spp. Zone, the equivalent of the upper
part of the standard ammonite Zone of Watinoceras
coloradoense (Kennedy 1984; Wright et al. in
Wright and Kennedy 1984) or Watinoceras spp.
(Hancock 1991) (see Nagm et al. 2010b for further
details).

1994. Neoptychites cephalotus (Courtiller); Chancellor et

al., 70, pl. 16, figs 19; pl. 17, figs 15; pl. 18, figs
13; pl. 26, figs 24. [with synonymy]
2008. Neoptychites cephalotus (Courtiller); Kennedy et al.,
159, pl. 3, figs 68; pl. 6, figs 15.
2010a. Neoptychites cephalotus (Courtiller); Nagm et al.,
489, figs 11GH, 12AB, 13AB.

MATERIAL: Three specimens (AfK 313, AUEN 16

17) from the upper part of the Galala Formation at the
north Wadi Qena section, co-occurring with Choffaticeras segne (Text-fig. 2).
MEASUREMENTS:
Specimen
D
Wb
Wh Wb/Wh U
AfK 313 63.3 (100) 42.9 (67.7) 32.9 (51.9) 1.30 5.1 (8.0)

DESCRIPTION: All three specimens are characterized

by very involute coiling with a small and shallow umbilicus (U ~8% of the diameter). The greatest whorl
breadth is situated shortly above the umbilical shoulder. Whorls are generally depressed, with a subtrigonal section (Text-fig. 11). The flanks are strongly convex, converging to a rounded venter. Ornamentation
and suture line are not seen due to the poor preservation.

Genus Neoptychites Kossmat, 1895

TyPE SPECIES: Ammonites telinga Stoliczka, 1865;
by the subsequent designation of Solger 1904 (= Ammonites cephalotus Courtiller 1860).
Neoptychites cephalotus (Courtiller, 1860)
(Text-figs 11, 13B)
1860. Ammonites cephalotus Courtiller, 248, pl. 2, figs 14.
1907. Neoptychites cephalotus (Courtiller); Pervinquire,
393, pl. 27, figs 14, text-fig. 152.

Text-fig. 11. Whorl section of Neoptychites cephalotus (Courtiller, 1860),

Lower Turonian of the Galala Formation, specimen AfK 313

75
CENOMANIAN TURONIAN AMMONITES FROM EGyPT

REMARKS: Large collections of Neoptychites

cephalotus have been described from Tunisia (Pervinquire 1907; Chancellor et al. 1994), France (Kennedy
and Wright 1979), and New Mexico, USA (Cobban and
Hook 1983). The last authors demonstrated a wide
range of intraspecific variation and dimorphism in the
species. This variation has also been documented in
Egyptian material by Nagm et al. (2010a). The present
material falls within the range of variation described in
the literature; it is close to the Tunisian material (Chancellor et al. 1994) and material described recently from
Morocco (Kennedy et al. 2008).
OCCURRENCE: Known from France, northern Spain,
Morocco, Algeria, Tunisia, Egypt, Israel, Syria, Jordan,
Cameroon, Madagascar, southern India, Colorado,
New Mexico (USA), northern Mexico, Trinidad,
Venezuela, Colombia, Brazil, Niger and Nigeria. In the
study area, this species co-occurs with Choffaticeras
segne and Thomasites rollandi which are dominant in
the Lower Turonian of Egypt (Nagm et al. 2010a, b).
It ranges into the basal Middle Turonian Collignoniceras woollgari Zone (Kennedy et al. 2008).
Family Pseudotissotiidae Hyatt, 1903
Subfamily Pseudotissotiinae Hyatt, 1903
Genus Thomasites Pervinquire, 1907
TyPE SPECIES: Pachydiscus rollandi Thomas and
Peron, 1889; by subsequent designation of Diener,
1925.
Thomasites gongilensis (Woods, 1911)
(Text-figs 12, 13C)

1911. Vascoceras gongilensis Woods, 282, pl. 21, fig. 7; pl.

22, fig. 1.
1981. Thomasites gongilensis (Woods); Wright and
Kennedy, 100, pl. 24, fig. 1; pl. 25, fig. 1; including varieties tectiformis Barber 1957, and lautus Barber
1957. [with full synonymy]
1989. Thomasites gongilensis (Woods); Meister, 38, pl. 16,
figs 35; pl. 17, figs 16; pl. 18, figs 13; pl. 19, figs
15; pl. 20, figs 15; pl. 21, figs 13.
2003. Thomasites gongilensis (Woods); Kennedy et al., 12,
pl. 5, figs 13.

MATERIAL: One specimen (AfK 314) from the upper

part of the Galala Formation at the north Wadi Qena
section, co-occuring with Choffaticeras segne (Textfig. 2).
MEASUREMENTS:
Specimen
D
Wb
Wh
Wb/Wh
U
AfK 314 81.8 (100) 42.3 (51.7) 43.5 (53.1) 0.97 19.3 (23.5)

DESCRIPTION: The specimen has a preserved diameter of ~82 mm and a moderately wide, deep umbilicus with broadly rounded umbilical shoulders. The
whorl section (Text-fig. 12) is only slightly compressed
(Wb/Wh = 0.97), with the greatest width at the umbilical shoulders. The flanks are convex, ending at
rounded ventrolateral shoulders. The venter is fairly
broad and flat. The suture line (Text-fig. 12) shows narrow lobes and broad, weakly incised saddles. Ornamental features are not preserved.
REMARKS: Wright and Kennedy (1981) revised
the genus Thomasites and discussed the relationship
of the present species to other species referred to the
genus. West African faunas of T. gongilensis (e.g.,

Text-fig. 12. Whorl section and suture line of Thomasites gongilensis (Woods, 1911), Lower Turonian of the Galala Formation, specimen AfK 314

76

EMAD NAGM AND MARKUS WILMSEN

Text-fig. 13. A Vascoceras kossmati Choffat, 1898 (A1, A2 - lateral and ventral views of AfK 311, 1). B Neoptychites cephalotus (Courtiller, 1860) (B1, B2
- apertural and ventral views of AfK 313, 1). C Thomasites gongilensis (Woods, 1911) (C1, C2 - apertural and lateral views of AfK 314, 1). D Thomasites
rollandi (Thomas and Peron, 1889) (D1, D2 - apertural and lateral views of AfK 316, 1). All specimens from the Lower Turonian of the Galala Formation

77
CENOMANIAN TURONIAN AMMONITES FROM EGyPT

Barber 1957) have been compared with Tunisian

material of T. rollandi by Chancellor et al. (1994).
They noted that T. gongilensis rarely shows coarse
umbilical tubercles (in contrast to juveniles of T.
rollandi), and generally has a less inflated, less triangular whorl section as well as a simpler suture line.
The present specimen of T. gongilensis differs from
T. rollandi in a wider umbilicus and a simple suture.
The Sinai specimen of T. gongilensis of El-Hedeny
(2002, fig. 4d, e) most probably represents Vascoceras cauvini.
OCCURRENCE: Thomasites gongilensis (Woods
1911) is common in the Lower Turonian of West
Africa. Kennedy et al. (2003) recorded it from the
Upper Cenomanian of France. In the study area, it
has been recorded from the Lower Turonian Choffaticeras spp. Zone, which is (see Nagm et al.
2010b) equivalent to the upper part of the standard
ammonite Zone of Watinoceras coloradoense
(Kennedy 1984; Wright et al. in Wright and
Kennedy 1984) or Watinoceras spp. (Hancock
1991). This is the first unequivocal record of T.
gongilensis from Egypt.
Thomasites rollandi (Thomas and Peron, 1889)
(Text-figs 13D, 14, 16A)
1889. Pachydiscus Rollandi Thomas and Peron, 25, pl. 17,
figs 13.
1994. Thomasites rollandi (Thomas and Peron); Chancellor et al., 75, pl. 19, figs 12; pl. 20, figs 112; pl. 21,
figs 19; pl. 22, figs 16; pl. 23, figs 19; text-fig.
14af. [with synonymy]
2010a. Thomasites rollandi (Thomas and Peron); Nagm et
al., 493, figs 13EG, 14AD, 15AD.

MATERIAL: Seven specimens (AfK 315, 316; AUEN

1822) from the upper part of the Galala Formation at the
Wadi El-Burga section, co-occuring with Choffaticeras
segne (Text-fig. 2).
MEASUREMENTS:
Specimen
D
Wb
Wh
Wb/Wh
U
AfK 315 88.7 (100) 47.1 (53.1) 50.2 (56.5) 0.93 10.3 (11.6)
AfK 316 88.9 (100) 49.6 (55.7) 51.1 (57.4) 0.97 9.3 (10.4)

DESCRIPTION: The incompletely preserved internal

moulds are smooth and characterized by involute coiling (U ~10% of diameter). The umbilical wall is high,
subvertical and ends at narrowly rounded umbilical
shoulders. The greatest whorl breadth is at the umbilical shoulder. The flanks are generally weakly convex
and converge strongly to a narrowly rounded venter.
The whorl section is compressed to slightly depressed
and shows a trigonal (Text-fig. 14) to ovoidal shape
(specimen AfK 316; Text-fig. 13D). The suture line
(Text-fig. 14) is characterized by four saddles, the one
between the external and adventive lobes being high,
broad and strongly incised.
REMARKS: Thomasites rollandi has been extensively
revised by Chancellor et al. (1994). They clarified the
range of its intraspecific variation and discussed its relationship to other species of the genus. Into synonymy of Th. rollandi they put numerous varieties
and species of Pervinquire (1907) and Douvill
(1928). The preservation of the specimens studied is
not good but they clearly belong to T. rollandi, especially specimen AfK 316 (Text-fig. 13D), which is
very close to the holotype as re-figured in Chancellor
et al. (1994, pl. 19).
OCCURRENCE: The species is well known from

Text-fig. 14. Whorl section and suture line of Thomasites rollandi (Thomas and Peron, 1889), Lower Turonian of the Galala Formation, specimen AfK 315

78

EMAD NAGM AND MARKUS WILMSEN

the Lower Turonian of Morocco, Tunisia, Algeria,

Egypt, Israel, Jordan, Syria, Lebanon, France and
Spain. It may also occur in England, Colombia and
Tadzhikistan (Chancellor et al. 1994). In the study
area, T. rollandi co-occurs with Lower Turonian
Choffaticeras segne, the range of which equates to
the upper part of the Watinoceras coloradoense
Zone of Kennedy (1984) and Wright et al. in Wright
and Kennedy (1984) (see Nagm et al. 2010b for details).
Genus Pseudotissotia Peron, 1897
TyPE SPECIES: Ammonites galliennei dOrbigny,
1850; by subsequent designation of Pervinquire,
1907.
Pseudotissotia nigeriensis (Woods, 1911)
(Text-figs 15, 16B)
1911. Hoplitoides nigeriensis Woods, 284, pl. 23, fig. 3; pl.
24, fig. 15; text-fig. 1.
1989. Pseudotissotia nigeriensis (Woods); Meister, 44,
pl. 21, figs 46; pl. 22, figs 14; pl. 23, figs 15; pl.
24, fig. 1; pl. 25, figs 17; pl. 26, figs 13; pl. 27, figs
13.
2005. Pseudotissotia nigeriensis (Woods); Gale et al., 186,
figs 10DE, 12AB, E, 13AB.

MATERIAL: Eleven specimens (AfK 328338) from

the uppermost part of the Galala Formation, in both
sections studied (~110 m above the base of the formation; Text-fig. 2).

MEASUREMENTS:
Specimen
D
AfK 328 68.5 (100)
AfK 329 91.1 (100)
AfK 330 84.0 (100)
AfK 331 85.8 (100)
AfK 332 78.5 (100)

Wb
19.7 (28.7)
33.4 (36.6)
20.1 (23.9)
27.3 (31.1)
23.4 (29.8)

Wh Wb/Wh
U
37.5 (54.7) 0.36 6.0 (8.7)
50.3 (55.2) 0.66 11.7 (12.8)
43.0 (51.1) 0.46 9.8 (11.6)
47.0 (54.7) 0.58 7.7 (9.0)
42.3 (53.8) 0.55 8.2 (10.4)

DESCRIPTION: All specimens are smooth, fully septate internal moulds with a maximum preserved diameter of 91 mm (specimen AfK 329, Text-fig. 16B46).
Coiling is involute (U ~10%) with a moderately deep
umbilicus and feebly rounded umbilical walls. The
flanks are high, subparallel, with convex inner and
converging outer parts. Two prominent keels at the
ventrolateral shoulders mark the contact with the venter, which is characterized by a siphonal keel, resulting
in the distinctive tricarinate appearance of the ventral
area (Text-fig. 15). The three keels have the same relative height in all specimens. The whorl section (Textfig. 15) is compressed, with the maximum breadth directly above the umbilical shoulder. The suture line
(Text-fig. 15) is characterized by very broad saddles and
narrow lobes; the adventive lobe is deeply incised.
REMARKS: The flat, tricarinate venter, the compressed whorl section and a suture line with broad
saddles and narrow lobes (deeply incised adventive
lobe) are characteristic of Pseudotissotia nigeriensis
(e.g., Meister 1989). Based on these characters, the
specimens from the upper Galala Formation of Wadi
Qena are unequivocally attributed to Woods species
and represent its first proven occurrence in Egypt.
Our specimens closely match the Nigerian material as
described by Woods (1911) and Meister (1989).

Text-fig. 15. Whorl section and suture line of Pseudotissotia nigeriensis (Woods, 1911), Lower Turonian of the Galala Formation, specimen AfK 328

79
CENOMANIAN TURONIAN AMMONITES FROM EGyPT

Text-fig. 16. A Thomasites rollandi (Thomas and Peron, 1889) (A1, A2 lateral and apertural views of AfK 315, 1). B Pseudotissotia nigeriensis (Woods, 1911) (B1,
B2 - lateral and apertural views of AfK 328, 1; B3 - lateral view of AfK 334, 1; B4, B5, B6 - apertural, lateral and ventral views of AfK 329, 1). All specimens from
the Lower Turonian of the Galala Formation

80

EMAD NAGM AND MARKUS WILMSEN

OCCURRENCE: The species is known from the

Lower Turonian of Nigeria, Algeria, Israel, northern
Mexico and Brazil. In the study area, it occurs in the
Lower Turonian, above the mid-Lower Turonian Choffaticeras spp. Zone. The P. nigeriensis horizon is thus
considered to be equivalent to the lower part of the
standard ammonite Zone of Mammites nodosoides (cf.
Kennedy 1984; Wright et al. in Wright and Kennedy
1984; Hancock 1991), since the last appearance of
Choffaticeras in Egypt is very close to the interpolated
first occurrence of Mammites nodosoides (Nagm et al.
2010b; see Text-fig. 22 and discussion below).
Genus Choffaticeras Hyatt, 1903
TyPE SPECIES: Pseudotissotia meslei Peron, 1897,
by original designation.

(102 m above the base of the formation) at the North

Wadi Qena section (Text-fig. 2).
DESCRIPTION: The specimens are large and characterized by a wide and fairly deep umbilicus, with vertical walls and a sharp umbilical shoulder. The flanks
converge to a narrow venter with a strong median, obtuse keel. The whorl section (Text-fig. 17) is compressed, lanceolate in shape and with the maximum
breadth at the umbilical shoulder. Ornament and suture
line are not preserved.
REMARKS: Freund and Raab (1969) described a large
collection (49 specimens) of Choffaticeras securiforme
from Israel, and discussed its relationship to C. (Leoniceras) luciae (Pervinquire) and C. (L.) massipianum
(Pervinquire). Although the specimens studied are very
poorly preserved, their whorl section (Text-fig. 17) and
the narrow venter, with a conspicuous median obtuse
keel, allow their safe assignment to C. securiforme.

Subgenus Choffaticeras Hyatt, 1903

Choffaticeras (Choffaticeras) securiforme (Eck, 1909)
(Text-fig. 17)
1909. Tissotia securiformis Eck, 187, figs 913.
1969. Choffaticeras securiforme (Eck); Freund and Raab,
60, pl. 9, fig. 1; text-fig. 11ik.
1996. Choffaticeras gr. securiforme (Eck); Meister and Abdallah, 15, pl. 9, fig. 1; text-fig. 6g.
2009. Choffaticeras (Choffaticeras) securiforme (Eck);
Nagm, 91, text-figs 5.19, 5.20A, 5.23H, I.

MATERIAL: Two fragmentary specimens (AUEN

2324) from the upper part of the Galala Formation

Text-fig. 17. Whorl section of Choffaticeras (Choffaticeras) securiforme

(Eck, 1909), Lower Turonian of the Galala Formation, specimen AUEN 23
from north Wadi Qena Section

OCCURRENCE: The species is known from the

Lower Turonian of Israel, Egypt and Tunisia. In the
study area, it comes from the middle Lower Turonian
Choffaticeras spp. Zone, which is (as defined by Nagm
et al. 2010b), equivalent to the upper part of the standard ammonite Zone of Watinoceras coloradoense
(Kennedy 1984; Wright et al. in Wright and Kennedy
1984) or Watinoceras spp. (Hancock 1991).
Choffaticeras (Choffaticeras) segne (Solger, 1903)
(Text-figs 18, 19)
1903. Pseudotissotia segnis Solger, 77, pl. 4, figs 1, 2; textfigs 1621.
1904. Schloenbachia quaasi Peron, 255, pl. 1, figs 13.
2007. Choffaticeras (Choffaticeras) quaasi (Peron); Barroso-Barcenilla and Goy, 464, figs 4(4,5), 5(1).
2007. Choffaticeras (Choffaticeras) segne (Solger); Barroso-Barcenilla and Goy, 468, figs 5(5), 6(13).
2008. Choffaticeras (Choffaticeras) segne (Solger); Kennedy et al., 162, fig. 9; pl. 3, figs 4, 5; pl. 7, figs 1, 2;
pl. 8, figs 16.
2009. Choffaticeras (Choffaticeras) quaasi (Peron); Nagm,
88, text-figs 5.18A, 5.23C.
2009. Choffaticeras (Choffaticeras) segne (Solger); Nagm,
94, text-figs 5.17B, 5.18C, 5.23EG.

MATERIAL: Twenty-seven specimens (AfK 317

327, AUEN 2533) from the upper part of the Galala
Formation (103104 m above the base of the formation) in both studied sections (cf. Text-fig. 2).

81
CENOMANIAN TURONIAN AMMONITES FROM EGyPT

MEASUREMENTS:
Specimen
D
AfK 317 101.3 (100)
AfK 318 107.4 (100)
AfK 319 114.1 (100)
AfK 320 81.8 (100)
AfK 321 57.9 (100)
AfK 322 45.5 (100)
AfK 323 43.8 (100)

Wb
36.8 (36.3)
31.8 (29.6)
36.9 (31.9)
22.5 (27.5)
20.0 (34.5)
15.4 (33.8)
12.9 (29.4)

Wh
Wb/Wh
U
50.2 (49.5) 0.73 13.0 (12.8)
47.5 (44.2) 0.66 17.4 (16.2)
67.0 (56.2) 0.55 23.1 (20.2)
42.9 (52.4) 0.52 10.0 (12.2)
33.7 (58.2) 0.59 3.6 (6.2)
25.9 (56.9) 0.59 5.8 (12.7)
23.8 (54.3) 0.54 5.4 (12.3)

DESCRIPTION: Both small and large specimens are

characterized by moderately involute coiling. The umbilicus has outward-inclined walls and rounded umbilical shoulders. The flanks are high and generally
convex, especially on the inner parts. Early whorls are
commonly ornamented by simple ribs and very weak
ventrolateral tubercles that disappear during ontogeny.
The venter is tricarinate in juvenile and sharp in adult
stages. The whorl section is compressed, with an oxycone shape (Text-fig. 18) and the greatest breadth on
the inner part of the flanks. The suture line is well preserved in all specimens and shows some variability
(Text-fig. 18). However, the sutures are generally characterized by broad and high saddles with a deep and
broad adventive lobe.

REMARKS: Barroso-Barcenilla and Goy (2007) described several species of Choffaticeras from Spain
and discussed their relationship. The Lower Turonian
of Egypt yielded abundant Choffaticeras (cf. Nagm
2009; Nagm et al. 2010a, b); however, only two
species are represented in Wadi Qena, i.e., Choffaticeras (Choffaticeras) securiforme and Choffaticeras
(C.) segne (and eight species are known in contemporaneous strata of Wadi Araba in the northern part of the
Eastern Desert; Nagm 2009; Wilmsen and Nagm, in
prep.). The well-preserved and large population of C.
(C.) segne from the Eastern Desert confirms the view
of Kennedy et al. (2008) that C. (C.) quaasi is a junior synonym of C. (C.) segne (Wilmsen and Nagm, in
prep.).
OCCURRENCE: The species is known from the
Lower Turonian of Tunisia, Israel, Syria, Jordan,
France and Spain. In the study area, it characterizes the
middle Lower Turonian Choffaticeras spp. Zone,
which is (as defined by Nagm et al. 2010b) equivalent
to the upper part of the standard Watinoceras coloradoense Zone (Kennedy 1984; Wright et al. in Wright
and Kennedy 1984) or the Watinoceras spp. Zone
(Hancock 1991).

Text-fig. 18. Whorl section and suture line of Choffaticeras (Choffaticeras) segne (Solger, 1903), Lower Turonian of the Galala Formation, A, B - specimen AfK 321;
C - specimen AfK 317; D - specimen AfK 323

82

EMAD NAGM AND MARKUS WILMSEN

Text-fig. 19. Choffaticeras (Choffaticeras) segne (Solger, 1903) from the Lower Turonian of the Galala Formation. A1, A2 - lateral and apertural views of AfK 319, 1;
A3, A4 - apertural and lateral views of AfK 317 , 1; A5, A6 - apertural and lateral views of AfK 325, 1

83
CENOMANIAN TURONIAN AMMONITES FROM EGyPT

Family Coilopoceratidae Hyatt, 1903

Genus Coilopoceras Hyatt, 1903
TyPE SPECIES: Coilopoceras colleti Hyatt, 1903, by
original designation.
Coilopoceras requienianum (dOrbigny, 1841)
(Text-figs 20, 21)
1841. Ammonites requienianum dOrbigny, 315, pl. 93,
figs 14.
1984. Coilopoceras requienianum (dOrbigny); Kennedy
and Wright, 282, pls. 35, 36, text-figs 15. [with
synonymy]
1989. Coilopoceras requienianum (dOrbigny); Luger
and Grschke, 388, pl. 46, text-figs 6a, e, 11, 12,
13ac.
2010a. Coilopoceras requienianum (dOrbigny); Nagm et
al., 495, figs 16AC, 17, 18AC.

MATERIAL: Fifteen specimens (AfK 339341,

AUEN 3445) from the upper part of the Umm
Omeiyid Formation in both sections studied (Textfig. 2).
MEASUREMENTS:
Specimen
D
Wb
Wh Wb/Wh
U
AfK 339 93.9 (100) 31.0 (33.0) 61.7 (65.7) 0.50 5.8 (6.1)
AfK 340 95.6 (100) 36.0 (37.6) 63.1 (66.0) 0.57 5.9 (6.1)
AfK 341 100.7 (100) 36.4 (36.1) 59.5 (59.0) 0.61 4.9 (4.8)

DESCRIPTION: All specimens are characterized by

very involute coiling. The umbilicus is small (U ~5
6%) and narrow, with low and rounded shoulders. The
flanks are convex and converge towards a pointed
venter. The whorl section (Text-fig. 20) is compressed,
with a lanceolate shape and the maximum breadth on
the inner flanks. Most specimens are smooth but some
of the small specimens have a few weak ribs on the inner whorls. The suture line (Text-fig. 20) shows a
broad and shallow external lobe, and a very broad,
open and bifid adventive lobe.
REMARKS: Cobban and Hook (1980) demonstrated
that many of the described species of the genus
Coilopoceras are probably synonyms, often being
based on small differences in the suture lines. Their
view is supported by the re-description of the type material of C. requienianum by Kennedy and Wright
(1984), who suggested the existence of sexual dimorphism in C. requienianum shown by smooth and ribbed
forms. The specimens from Wadi Qena closely correspond to the type material. However, some are more inflated and the smooth morphotype predominates.
OCCURRENCE: Upper Turonian of Madagascar,
Egypt, North and South America, France and Spain. In
the study area, C. requienianum characterizes the lower
Upper Turonian zone of Coilopoceras requienianum,
which is (as defined by Nagm et al. 2010b) equivalent
to the lower part of the standard ammonite Zone of
Subprionocyclus neptuni (Hancock 1991).

Text-fig. 20. Suture line and whorl section of Coilopoceras requienianum (dOrbigny, 1841), Upper Turonian of the Umm Omeiyid Formation, specimen AfK 339

84

EMAD NAGM AND MARKUS WILMSEN

Text-fig. 21. Coilopoceras requienianum (dOrbigny, 1841) from the Upper Turonian of the Umm Omeiyid Formation. A1, A2 - lateral and apertural views of AfK 339, 1;
A3, A4 - apertural and lateral views of AfK 341, 1

85
CENOMANIAN TURONIAN AMMONITES FROM EGyPT

BIOSTRATIGRAPHICAL AND PALAEOBIOGEOGRAPHICAL REMARKS

The first marine transgression in the study area took
place during the Late Cenomanian, documented by the
deposition of the shallow marine Galala Formation
above the Palaeozoic Naqus Formation, with a prominent sequence boundary (major non-sequence) at the
base of the Galala Formation. The early Late Cenomanian is known as a time of major eustatic sea-level
rise (e.g., Wilmsen 2003; Wilmsen et al. 2005). The
lower part of the Galala Formation is characterized by
glauconitic, silty sediments, rich in oysters distinctive of
Upper Cenomanian successions elsewhere in Egypt
(Nagm 2009). Approximately 35 m above the base of
the formation, the sediments have yielded the early
Late Cenomanian ammonite Neolobites vibrayeanus
(see Wiese and Schulze 2005 for details of the species).
It defines the oldest ammonite zone in the study area
(Text-fig. 2), equivalent to the northwest European standard ammonite Zone of Calycoceras (Proeucalycoceras) guerangeri (Kennedy 1984; Wright et al. in
Wright and Kennedy 1984; Hancock 1991) and the
Tethyan standard ammonite Zone of Calycoceras naviculare (Text-fig. 22; see Nagm et al. 2010b for full details). The beds of the Galala Formation below the first
occurrence of N. vibrayeanus are included in the N. vibrayeanus Zone (cf. Nagm 2009; Nagm et al. 2010b).
N. vibrayeanus ranges for about 35 m above its first occurrence, defining the top of the zone (Text-fig. 2). It is
followed by an ammonite-barren 2025 m interval rich
in the oyster Costagyra olisiponensis. Based on the latest Cenomanian age of the overlying strata, the barren
interval is regarded here as equivalent to the middle Upper Cenomanian Metoicoceras geslinianum Zone of
Kennedy (1984), Wright et al. in Wright and Kennedy
(1984) and Hancock (1991; see Text-fig. 22).
Above the barren interval, ammonites are common again. The lowest eight metres are characterized
by high carbonate contents and yield, in the lower
part, Vascoceras cauvini. This is the index of the
eponymous uppermost Cenomanian ammonite zone in
Egypt, largely equivalent to the Neocardioceras juddii
Zone of Kennedy (1984), Wright et al. in Wright and
Kennedy (1984) and Hancock (1991), in the Cenomanian standard ammonite zonation (Nagm et al.
2010b). Euomphaloceras costatum has been recorded
from the upper part of the Vascoceras cauvini Zone, for
the first time from Egypt. This species co-occurs with
Neocardioceras juddii in the United States (Cobban et
al. 1989). In the study area, the upper limit of this zone
is considered to mark the CenomanianTuronian (C
T) boundary (cf. Nagm et al. 2010b).

Text-fig. 22. Ammonite biostratigraphy of the Upper Cenomanian and Lower

Turonian. Vertically hatched fields represent stratigraphic gaps. See text for
further explanations

Two metres above the inferred CT boundary (a

short distance below the 100-m-level above the base of
the Galala Formation in both sections; Text-fig. 2),
Early Turonian vascoceratids (V. globosum globosum
and V. kossmati) become abundant in high-carbonate
sediments. The absence of Vascoceras proprium and
Pseudaspidoceras flexuosum suggests a stratigraphic
gap at this level (see Text-fig. 22), comparable to the
situation in some sections of Wadi Araba in the northern part of the Eastern Desert (Nagm et al. 2010b).
Two horizons with representatives of the genus Choffaticeras have been recognized directly above the lowermost Turonian vascoceratid interval (Text-fig. 2): the
lower one with Choffaticeras (Choffaticeras) securiforme; and the upper one with C. (C.) segne. The ~7 m
thick Choffaticeras interval characterizes the middle
Lower Turonian (Nagm et al. 2010b), equivalent to the
upper part of the standard ammonite Zone of
Watinoceras devonense (Kennedy 1984; Hancock
1991; see Text-fig. 22). This part is followed by the
highly fossiliferous uppermost interval of the Galala
Formation, yielding typical Nigerian ammonites, such
as Pseudotissotia nigeriensis (Text-fig. 2), dated as late
Early Turonian (Meister 1989). In agreement with this
observation, Nagm et al. (2010b) proposed that the last
appearance of the genus Choffaticeras approximates
the first appearance of the index of the standard upper
Lower Turonian Mammites nodosoides ammonite
Zone of Kennedy (1984) and Hancock (1991). Therefore, the P. nigeriensis Zone (or horizon) seems to be

86

EMAD NAGM AND MARKUS WILMSEN

equivalent to the lower part of the M. nodosoides

Zone. The upper boundary of this zone is marked by
a prominent unconformity in the LowerMiddle Turonian boundary interval (Text-fig. 2). The same unconformity has been recognized in the sections of
Wadi Araba in the northern part of the Eastern Desert;
Nagm et al. 2010b), and it also has its equivalent in
shallow shelf settings of the northern Tethys margin
(see Niebuhr et al. 2011). This unconformity also represents the contact of the Galala and Umm Omeiyid
formations. The lower part of Umm Omeiyid Formation is dominated by siliciclastics and is barren of
macrofauna. In the upper part of the formation, fossiliferous strata yielded the early Late Turonian ammonite Coilopoceras requienianum, the zone of which
is (as defined by Nagm et al. 2010b) equivalent to the
lower part of the Upper Turonian standard ammonite
Zone of Subprionocyclus neptuni (Geinitz) (Kennedy
1984; Hancock 1991; see Text-figs 2, 22). This observation suggests that the unfossiliferous part of the
lower Umm Omeiyid Formation corresponds, in a
broad sense, to the Middle Turonian. Interestingly,
also in the sections of Wadi Araba, Middle Turonian
strata are barren of macrofossils (Nagm 2009; Nagm
et al. 2010a, b).
The early Late Cretaceous eustatic transgression,
with a significant sea-level rise that peaked during the
Late Cenomanian and Early Turonian, has been well
documented (Hancock and Kauffman 1979; Hardenbol
et al. 1998; Sharland et al. 2001; Wilmsen 2003; Wilmsen et al. 2005, 2010). Rising sea level may be a
straightforward explanation for the wide geographic
distribution of many CenomanianTuronian ammonites (Wiedmann 1988; Lehmann and Herbig 2009;
Nagm et al. 2010a). The palaeontological record documents ammonite geographic expansion during the
Late Cenomanian, expressed by the presence of a single large Tethyan province, followed by biogeographic
differentiation during the Early Turonian, expressed by
Boreal, Vascoceratid and Puzosiid provinces (Wiedmann 1988; Lehmann and Herbig 2009). The palaeobiogeographic affinities of the studied fauna correspond to this pattern (see Nagm et al. 2010a for data
from Wadi Araba). The early and mid-Late Cenomanian ammonite fauna is biogeographically not very diagnostic. The latest Cenomanian incipient provincialism is documented by the presence of the typically
North and West African Vascoceras cauvini. The Early
Turonian assemblage, studied herein, belongs to the
Tethyan Vascoceratid Province, with strong influences
of Nigerian faunas (Vascoceras globosum globosum,
Thomasites gongilensis, Pseudotissotia nigeriensis),
proving the faunal connection to Central and West

Africa via the Trans-Saharan Seaway (see Collignon

and Lefranc 1974; Meister et al. 1992). The palaeobiogeographic affinity to the Boreal Realm is much
less noticeable in the study area, with only 50% of the
ammonites of Wadi Qena known also from that Realm.
It is noteworthy that, in the contemporaneous successions of the northern part of the Eastern Desert (Wadi
Araba, ca. 130 km to the north), 85% of the Early Turonian ammonoids identified are also known from the
Boreal Realm (Nagm et al. 2010a). This observation
may indicate relatively steep northsouth environmental gradients.
Acknowledgements
We greatly acknowledge constructive reviews by F.
Barroso-Barcenilla (Madrid), C. Meister (Geneva) and an
anonymous referee. Furthermore, we thank W.J. Kennedy
(Oxford) for the discussion of taxonomic problems as well
as I. Walaszczyk (Warszawa) and C.J. Wood (Minehead) for
their editorial work. R. Winkler and M. Rthel (Senckenberg
Dresden) did the photographic and collection work at
SNSD. The financial support of DAAD for the stay of EN
in Germany is greatly acknowledged. This paper is a contribution to German Research Foundation (DFG) project WI
1743/6-1.

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