Palynology

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Early Cretaceous palynostratigraphy of the

Abu Tunis 1x borehole, northern Western
Desert, Egypt, with emphasis on the possible
palaeoclimatic effect upon the range of
Dicheiropollis etruscus in North Africa

Amr S. Deaf, Ian C. Harding & John E.A. Marshall

To cite this article: Amr S. Deaf, Ian C. Harding & John E.A. Marshall (2016) Early Cretaceous
palynostratigraphy of the Abu Tunis 1x borehole, northern Western Desert, Egypt, with
emphasis on the possible palaeoclimatic effect upon the range of Dicheiropollis etruscus in
North Africa, Palynology, 40:1, 25-53, DOI: 10.1080/01916122.2014.993480

To link to this article: http://dx.doi.org/10.1080/01916122.2014.993480

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 Palynology, 2016
Vol. 40, No. 1, 25
53, http://dx.doi.org/10.1080/01916122.2014.993480

Early Cretaceous palynostratigraphy of the Abu Tunis 1x borehole, northern Western Desert,
Egypt, with emphasis on the possible palaeoclimatic effect upon the range of Dicheiropollis etruscus
in North Africa
a,b
Amr S. Deaf *, Ian C. Hardinga and John E.A. Marshalla
a
School of Ocean and Earth Science, National Oceanography Centre, University of Southampton, European Way, Southampton,
SO14 3ZH, UK; bGeology Department, Faculty of Science, Assiut University, Assiut, 71516, Egypt
Recent hydrocarbon exploration in the northern sector of the Western Desert in Egypt has revealed relatively rich
hydrocarbon accumulations, mainly of gas, and demonstrate promising future prospects. In order to improve our
understanding of this area and to provide a biostratigraphical framework for the hitherto poorly dated Lower
Cretaceous successions, a palynological analysis was carried out on 57 ditch cutting samples from the Abu Tunis 1x
borehole. Palynostratigraphic investigation on these samples has enabled the identification of three new
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palynostratigraphically defined age divisions with three corresponding palynozones defined by first uphole occurrences
of gymnosperm and angiosperm pollen and dinoflagellate cysts. Spore and pollen grains recovered from the Abu Tunis
1x borehole show the characteristics of the pre-Albian Dicheiropollis/Afropollis Phytogeographical Province.
Discrepancies in the reported range of Dicheiropollis etruscus, when compared with earlier (Berriasian) appearances in
West Africa and later (late Hauterivian) appearances in East Africa, may be attributed to palaeoecological factors.
Dicheiropollis etruscus is accepted as having a cheirolepidiacean conifer affinity and is regarded as having been
produced by a thermophilous plant. Here, we suggest that Dicheiropollis etruscus was adapted to arid conditions.
Dicheiropollis etruscus thus first appeared in hot, dry palaeo-subtropical African regions, but as Western Gondwana
broke up and the African Plate moved northeast during/after the Late Jurassic, the region that is now present-day
Egypt, Libya and Sudan had moved by the late Hauterivian into a subtropical position; the ensuing increased aridity
thus allowed Dicheiropollis etruscus to migrate into these areas.
Keywords: palynostratigraphy; Cretaceous; Western Desert; Egypt; Dicheiropollis etruscus; North Africa

1. Introduction successions in the northern Western Desert. In this

Recently, significant new hydrocarbon discoveries have
been documented in the northern sector of the Western
Desert of Egypt, the second most important oil-pro-
ducing area in Egypt. These new hydrocarbon discov-
eries are in Upper Jurassic and Lower Cretaceous
reservoirs. Therefore, in addition to the known Ala-
mein (Aptian), Bahariya (early Cenomanian) and Abu
Roash (late Cenomanian
Turonian) oil-producing
horizons, the lower part of the Lower Cretaceous suc-
cession (Neocomian
Barremian) is now the current
target for oil and gas explorations. However, the sedi-
ments of this geological interval (i.e. Lower Creta-
ceous) have not previously been subject to either
detailed biostratigraphical study or lithostratigraphical
correlation. Even the basic division of the different
time
rock units is low resolution, as operating compa-
nies label this part of the succession on their logs as ‘no
information’. For this reason, operators are now inter-
ested in large-scale and detailed biostratigraphical and
lithostratigraphical correlations on several Cretaceous
regard, Deaf et al. (2014) addressed similar middle
Upper Cretaceous biostratigraphical and lithostrati-
graphical problems of the Abu Tunis 1x borehole, and
identified quantitatively a mid Albian
early Cenoma-
nian Afropollis jardinus ‘acme’ as an important regional
biostratigraphical marker in the mid Cretaceous.
The Lower Cretaceous sediments of the northern
basinal area of Egypt were mainly deposited in very
shallow marine (brackish to coastal) to inner neritic
open marine conditions (Kerdany & Cherif 1990; Said
1990), which were unfavourable for the proliferation of
planktonic forams and calcareous nannofossils. Conse-
quently, no independent age control is available for
these successions. However, we have determined the
ages we assign to our material by correlation with the
known ranges of index palynological taxa recorded
from successions within the same phytogeographical
province, which themselves have independent age con-
trol. The new zonal scheme we erect uses the same defi-
nitions used by Schrank & Ibrahim (1995) and Ibrahim

*Corresponding author. Email: amr.daif@science.au.edu.eg

Ó 2015 AASP
The Palynological Society

 26 A.S. Deaf et al.

& Schrank (1996) where possible to provide a unified
zonation scheme for the Early Cretaceous of the north-
ern Western Desert. Thus, this study aims to: (i) inte-
grate lithostratigraphical and biostratigraphical
scheme for the Abu Tunis 1x borehole sequence; (ii)
propose a unified informal zonal scheme for the Early
Cretaceous of the northern Western Desert; (iii) corre-
late biostratigraphical results with those in other palae-
ogeographically related areas, such as the African and
South American (ASA) Phytogeoprovince based on
fossil sporomorphs, and the Tethyan Realm for dino-
flagellate cysts; (iv) provide the first quantitative
description of the characteristic Egyptian pre-Albian
Dicheiropollis/Afropollis palynoflora; (v) provide a
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taxonomic study of all the stratigraphically and/or eco-
logically significant taxa recorded.
2. Geological settings and lithostratigraphical history
Said (1962) divided the Egyptian continental mass into
two main provinces. The Stable Shelf is represented by
the southern Western Desert bordering the Nubian
Shield and the Eastern Desert as far as the northern
eastern margin of the South Galala Plateau, and south-
ern Sinai (Figure 1). The Unstable Shelf lies to the
north of the Stable Shelf with a Western Subprovince
(i.e. the northern Western Desert) and an Eastern Sub-
province (i.e. northern Sinai), with the northern

Figure 1. Structural map of Egypt showing the boundary between the Stable and Unstable shelves, and distribution of different
Mesozoic tectonic elements (after Kerdany & Cherif 1990).
 Palynology
Eastern Desert defined as a transitional zone between
the Eastern and Western Subprovinces. The study area
(i.e. the Faghur area) is located at the far northwest of
the Western Subprovince (Figure 1). Topographically,
the Egyptian northern Western Desert area is charac-
terised by an almost plain surface, but with complex
subsurface structures hidden underneath Neogene sedi-
mentary cover. The northern Western Desert is a vast
area of about 250,000 km2 and is composed of a thick,
gently northward dipping sedimentary sequence, rang-
ing in age from Cambro
Ordovician to Recent. The
sequence attains its maximum thickness in the Abu
Gharadig Basin (8
9 km), while to the north it attains
only 3
6 km (Hantar 1990). Mesozoic rocks crop out
in southern Egypt and in northern Sinai, where an
almost-complete sequence from the Triassic to the Cre-
taceous has been recorded. However, in the northern
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Western Desert, Mesozoic rocks are buried beneath
younger Neogene sediments and are only known from
the subsurface (Kerdany & Cherif 1990).
Deposition of the middle-Upper Jurassic carbo-
nates of the northern Western Desert occurred during
a late Callovian acme of marine transgression when the
Masajid Formation was deposited. However, the
Egyptian northern basins witnessed a major latest
Jurassic regression related to the late Tithonian global
eustatic sea level fall (Kerdany & Cherif 1990; Guiraud
1998). As a result, the Upper Jurassic rocks (Masajid
Formation) east of the Faghur-Maamura high (the
area of the present study) and in the Umbarka and
Alamein basins were exposed and subjected to erosion,
forming a regional unconformity surface (Kerdany &
Cherif 1990). This unconformity was also recorded in
the Abu Gharadig and Gindi basins, about 200 km
and 380 km, respectively, to the southeast of the
Faghur-Maamura high. But in the extreme north, con-
tinuous sedimentation took place in the Matruh and
Sidi Barrani basins (Kerdany & Cherif 1990). This
major non-deposition and break in sedimentation con-
tinued until the early Neocomian because of the Late
Jurassic
early Neocomian (Neocimmerian) tectonic
event (Kerdany & Cherif 1990). This event is related to
the eastward movement of the African Plate with
respect to the European Plate as the Atlantic Ocean
opened (Meshref 1990; Bumby & Guiraud 2005),
which resulted in a regional uplift of some of the sedi-
mentary basins of the northern western part of Egypt
(Meshref 1990). Deposition of the Lower Cretaceous
sediments had started by the late Neocomian (early
Hauterivian), it was mainly affected by synsedimentary
tectonics and was also related to the movement of the
African Plate towards Laurasia
i.e. the opening of
the Atlantic Ocean
and to the Neotethyan rifting
(Meshref 1990; Guiraud et al. 2001). Deposition was
mainly controlled by a west
northwest folding trend
27
(Meshref 1990). In general, Lower Cretaceous sedi-
mentation exhibits a regressive character represented
by a clastic sequence comprising the Alam El Bueib
and Dahab formations, with the exception of the Ala-
mein Dolomite (Kerdany & Cherif 1990; Said 1990).
3. Material
The Abu Tunis 1x borehole was drilled in the Faghur
area (Lat. 31 160 0800 N, Long. 26 500 4100 E) by Western
Desert Operating Petroleum Company (WEPCO) in
(1968), which is located at the central Matruh Basin, to
the east of the Faghur-Maamura High, and north of
the Umbarka Subbasin (Figure 1). The sample material
investigated comprised 57 ditch-cutting samples from
3094
2240 m (10,150
7350 ft), which spans most of
the Lower Cretaceous sequence of the northern West-
ern Desert. A summary of the samples studied, their
depth, palynological status and total recovery of paly-
nomorphs in terms of grains/gram of sediments is given
in the online supplementary Appendix 1.
4. Methods
4.1. Palynological processing technique
The dominantly siliclastic samples were processed
using a standard palynological processing technique.
This comprised hydrochloric acid (HCl, 36%) pre-
treatment of crushed samples to remove carbonates
followed by addition of 60% hydrofluoric acid (HF) to
digest the silicates (e.g. Phipps & Playford 1984; Green
2001). An exotic spike of Lycopodium spores was
added to each sample after the first HF decanting for
absolute abundance analysis. Samples were sieved at
15 mm and two permanent slides were made using
Elvacite 2044 as a mounting medium. More details on
the technique are in Deaf et al. (2014). No ultrasonic
treatment or oxidation was employed in order to con-
duct unbiased palynofacies analyses. Slides, residues
and samples are stored in the Geological Museum,
Geology Department, Faculty of Science, Assiut Uni-
versity, Egypt.
4.2. Qualitative and quantitative palynological
investigations
Palynological residues yielded well-preserved and
diverse assemblages of sporomorphs with less frequent
occurrences of lower diversity dinoflagellate cysts.
Qualitative investigation was based on the light micro-
scope description of the palynological assemblages
using an Olympus (BX41) transmitted-light micro-
scope (serial no. 8B25715) equipped with an Infinity-1
digital camera used for photomicroscopy. The
 28 A.S. Deaf et al.

taxonomic identification of the spores, pollen and dino-

flagellate cysts was made to generic and specific levels
with reference to the original descriptions and diagnoses
of the species in question. More details of systematic
palynology are in the online supplementary Appendix 2.
Quantitative determinations were based on palyno-
logical counts against the number of out-of-count
Lycopodium spores recorded. The absolute abundance/
concentration (specimens/gram) of each fossil category
counted (i.e. spores, gymnosperm pollen, etc.) was
obtained by applying the absolute abundance formula
(1) of Stockmarr (1971) as follows:

Sc £Lt £t
c¼ (1)
Lc £w

where:
c D concentration D total number of specimens/
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gram dried sediment;

Sc D number of specimens counted;
Lt D number of Lycopodium spores/tablet;
t D number of tablets added to the sample;
Lc D number of Lycopodium spores counted;
w D weight of dried sediments (g).

Two hundred and fifty specimens were counted

from each preparation where possible, this number pro-
viding a total maximum error of 7% according to the
Stockmarr (1971) curve. Further scanning of the rest of
each slide was made to detect any rare, out-of-count
species. The abrupt cut-off of the downhole distribution
of most of the selected taxa proved the occurrence of
limited cavings in the studied borehole samples.

5. Palynostratigraphy
5.1. Preliminary notes
A literature appraisal of palynological work on Egyp-
tian Cretaceous material has shown that some of the
age assessments are equivocal either because of the use
of species ranges, which have no independent age cali-
bration, or due to inaccurate taxonomic identification
of species. For example, the work of Mahmoud and
Moawad (2000) on subsurface Upper Jurassic
lower
Upper Cretaceous deposits from the West Tiba-1 bore-
hole in the northern Western Desert provides impor-
tant information on the Upper Jurassic. However, for
the Lower Cretaceous, their assignment of Afropollis
jardinus to the Aptian is unjustified, because Afropollis
jardinus unequivocally marks the base of the Albian
(e.g. Doyle et al. 1982) in the Albian
Cenomanian
Elaterate Province. Thus, it is necessary to compile paly-
nological biostratigraphical data by selecting only those
index species with independently calibrated age ranges
(Figure 2). The age assignments for the succession in the
Figure 2. Compilation of the biostratigraphical ranges of
important Early Cretaceous marker species in different phyto-
geographical provinces of North and West Africa, northern
South America and the Tethyan Realm. Sources for African
ranges: Doyle et al. 1982 (8, 9); G€ubeli et al. 1984 (8); Hochuli
1981 (4, 8, 9); Schrank & Mahmoud 1998 (1, 3); Thusu & Van
Der Eem 1985 (1
7); Thusu et al. 1988 (1
7). Sources for
northern South American ranges: Regali & Viana 1989 (4,
6
9); Regali et al. 1974 (4, 7). Sources for Tethyan ranges:
Davey & Verdier 1973 (18); Davey & Verdier 1974 (18
20);
Duxbury 1983 (19); Erba et al. 1999 (10); Foucher et al. 1994
(17, 19, 20); Habib & Drugg 1983 (10, 11); Hoedemaeker &
Leereveld 1995 (11, 12, 17); Leereveld 1997a (11, 12); Leereveld
1997b (13
16); Lister & Batten 1988 (19, 20); Srivastava 1984
(15); Thusu et al. 1988 (10
13, 17); Torricelli 2000 (10, 11, 13,
14, 16, 18, 20); Torricelli 2001 (10, 13, 18
20); Torricelli 2006
(14, 18); Wilpshaar 1995 (11, 13, 15, 16).
 Palynology
Abu Tunis 1x borehole are thus based on diagnostic
palynomorph taxa correlated with better-dated contem-
poraneous regional and interregional palynofloral
assemblages, in the context of the pre-Albian Dicheiro-
pollis etruscus/Afropollis Phytogeographical Province of
Herngreen et al. (1996) for sporomorphs, and the
Tethyan Realm for the dinoflagellate cysts.
It is important to note that the micropalaeontologi-
cally calibrated palynological work of Thusu et al.
(1988) on Libyan Lower Cretaceous sediments, which
is in part correlated to the formal Lower Cretaceous
dinoflagellate zonation of the European Tethys (Leere-
veld 1997a), should be viewed with some caution. Leer-
eveld (1997a) pointed out that the lower part of
Miospore Association V of Thusu et al. (1988), of pro-
posed Berriasian age, contains the typical Valanginian
species Calpionellites darderi. The middle part of Mio-
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spore Association V, which Thusu et al. (1988)
assumed to be of Valanginian age, is actually consistent
with an Hauterivian age based on the presence of the
dinoflagellate index species Muderongia staurota (Leer-
eveld 1997a). This revised age assessment made by
Leereveld (1997a) for the Miospore Association V of
Thusu et al. (1988) will be followed here.
Moreover, the ranges of some of the pre-Aptian
palynomorphs of the palaeotropical region, e.g. Sene-
gal and the Ivory Coast, differ greatly from those
recorded in the palaeosubtropical region, e.g. Congo
and Gabon, for example the index gymnosperm pollen
Dicheiropollis etruscus (Jardine et al. 1974; Salard-Che-
boldaeff 1990). As there is no palynologically
Figure 3. Palaeogeographical map showing the position of north Egypt during Berriasian time (after Lawver et al. 2004).
29
independent biostratigraphical evidence, it is assumed
here that these regions contain at least a few ecologically
controlled species with different first appearances. As a
result, the pre-Aptian stratigraphical units of the palaeo-
tropical and palaeosubtropical regions could be biostrati-
graphically diachronous. From a palaeogeographic point
of view, northern Egypt was located at around 5 to 8 N
(Figure 3) during the Berriasian
Hauterivian (Lawver
et al. 2004, www.ig.utexas.edu/research/projects/plates/),
where Libya and Senegal were nearly confined to the
same palaeolatitude, where they possessed palynofloras
of more similar stratigraphic ranges than that of the
palaeosubtropical regions mentioned above.
Therefore, the micropalaeontologically dated paly-
nological work carried out on the pre-Aptian of Libya
(Thusu & van der Eem 1985; Thusu et al. 1988) and that
carried out on the Aptian of Senegal (Jardine &
Magloire 1965) will be employed here for dating the pre-
Aptian and Aptian sequences studied here, respectively.
Sporomorph ranges are used here for palynostrati-
graphical purposes because their vertical distribution is
continuous, and the angiosperm pollen grains in partic-
ular exhibit reliable lineage trends (e.g. the Afropollis
complex). The dinoflagellate cysts provide additional
supporting evidence for ages, but it should be noted
that some dinoflagellate cyst taxa exhibit facies control
in Lower Cretaceous sediments (Thusu et al. 1988).
For the proposed biozonation and age assessments of
the studied borehole samples, the vertical quantitative
distributions of all taxa recovered have been arranged
by their highest appearance data, i.e. tops (Figure 5) to
 30 A.S. Deaf et al.
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Figure 4. Palaeogeographic map showing the position of north Egypt during Aptian time (after Lawver et al. 2004).
exclude base range extension by cavings. The up-hole
first appearance datum (FAD) of the index spores and
pollen has been used in the delineation of most of the
biostratigraphical units, with their last appearance
datum (LAD) being used when no FAD is available,
and also to provide supporting evidence to the age
determinations. The numbers in parentheses after the
names of the taxa refer to the position of these species
in the quantitative range chart.
5.2. Palynozonation and age appraisal
5.2.1. Late Jurassic
Sample. Sample 1 (depth 10,150 ft/3094 m)
Associated palynoflora. Deltoidospora spp. (1),
Triplanosporites sp. (3), Deltoidospora halii (5), Conca-
vissimisporites punctatus (10), Concavisporites spp.
(11), Cicatricosisporites spp. (12), Deltoidospora minor
(23), Dictyophyllidites harrisii (25), Cibotiumspora jurie-
nensis (36), Deltoidospora australis (37), Balmeiopsis
limbatus (60), Araucariacites australis (63), Classopollis
spp. (64), Inaperturopollenites undulatus (65), Taxacites
sahariensis (71).
Remarks. The palynoflora of sample 1 is mainly domi-
nated by psilate pteridophyte spores and spherical gym-
nosperm pollen taxa (Figure 5; Plate 1). No angiosperm
pollen or dinoflagellate cysts have been recorded.
Age appraisal. This sample is characterised by
few long-ranging spores (e.g. Deltoidospora spp.,
Triplanosporites sp. and Dictyophyllidites spp.) and
gymnosperm pollen (e.g. Araucariacites australis, Bal-
meiopsis limbatus and Classopollis spp.) taxa, which
range in age from the Late Jurassic to the Early Creta-
ceous in Egypt and northeast Libya (Schrank 1984;
Thusu & van der Eem 1985; Thusu et al. 1988).
The lack of additional samples below 10,150 ft
(3094 m) does not permit a more precise age for this
sample. The species Impardecispora apiverrucata,
Aequitriradites spinulosus and Pilosisporites trichopapi-
losus, which are considered as early Berria-
sian
Valanginian marker forms (Schrank &
Mahmoud 1998; Thusu et al. 1988), and Dicheiropollis
etruscus, which is characteristic of the late Hauteri-
vian
early Barremian (Hochuli 1981; Thusu et al.
1988), are absent from sample 1.
The abrupt appearance of all of these forms in the
overlying sample 2 (10,100 ft/3078 m) is therefore com-
patible with a Jurassic age for sample 1 (10,150 ft/
3094 m). From a lithostratigraphical point of view, the
position of this sample below an unconformity con-
firms the major unconformity, which separates the
Jurassic from the overlying Cretaceous succession in
most of the northern Western Desert (Morgan 1990).
The Upper Jurassic (Kimmeridgian)
Lower Creta-
ceous (lower Neocomian) rocks were exposed and sub-
jected to erosion in most of the northwestern basinal
areas, except in the extreme north, where continuous
sedimentation took place in the Matruh and Sidi Bar-
rani basins (Kerdany & Cherif 1990). A late Jurassic
age is also in accordance with the age of the Masajid
Formation, where the drilling company (WEPCO
 Palynology 31
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Figure 5. Quantitative distribution chart (grains/gram) by highest appearance of the palynomorphs recovered from the Abu
Tunis 1x borehole.
 Downloaded by [Assiut University] at 08:01 24 February 2016

32

Figure 5. (Continued )
A.S. Deaf et al.
 Downloaded by [Assiut University] at 08:01 24 February 2016

Figure 5. (Continued )
Palynology
33
 34 A.S. Deaf et al.
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Figure 5. (Continued )
1968) originally designated this part of the borehole as
representing the Masajid, and also allocated a Jurassic
age (Figure 6).
5.2.2 Palynozone 1 (late Hauterivian
early
Barremian): Dicheiropollis etruscus Total
Range Zone
Samples. This zone includes samples 2 to 9, which
were recovered from a depth of 10,100 to 9750 ft
(3078
2972 m).
Definition. Total range of Dicheiropollis etruscus (76).
Associated palynoflora. Deltoidospora spp. (1), Conca-
vissimisporites punctatus (10), Concavisporites spp.
(11), Cicatricosisporites spp. (12), Dictyophyllidites har-
risii (25), Cibotiumspora jurienensis (36), Deltoidospora
australis (37), Trilobosporites hannonicus (51), Imparde-
cispora uralensis (58), Balmeiopsis limbatus (60), Class-
opollis classoides (61), Araucariacites australis (63),
Inaperturopollenites undulatus (65), Taxacites saharien-
sis (71), Cribroperidenium sp. (122), Circulodinium
 Palynology
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Figure 5. (Continued )
distinctum (125), Muderongia tomaszowensis (141),
Muderongia spp. (146) and Phoberocysta spp. (156).
Remarks. Samples 2
9 are similar to sample 1 in their
palynofloral content, where smooth-walled pterido-
phytic spores and gymnosperm pollen grains dominate
the microfossil assemblage (Figure 5), and there are
very rare occurrences of some dinoflagellate cysts.
Samples 2
9 witnesses the first appearance of several
of the index sporomorph species mentioned below.
Age appraisal. Samples 2
9 witness the first appear-
ance of the gymnosperm index pollen Dicheiropollis
etruscus, along with the marker spores Impardecispora
apiverrucata, Aequitriradites spinulosus and Pilosispor-
ites trichopapillosus (Figure 5; Plate 1, figures 14
17,
35
20, 21). Thusu & van der Eem (1985) and Thusu et al.
(1988) recorded Impardecispora apiverrucata from cal-
pionellid-dated Valanginian rocks in northeast Libya.
Aequitriradites spinulosus ranges from the calpionellid-
dated Valanginian rocks of Libya to the foraminifera-
dated early Cenomanian rocks of Egypt (Thusu & van
der Eem 1985; Thusu et al. 1988; Schrank & Ibrahim
1995). Schrank & Mahmoud (1998) regarded Imparde-
cispora apiverrucata and Aequitriradites spinulosus as
Berriasian
Valanginian marker species in Egypt,
based on well-dated European, Libyan and other
sequences. Pilosisporites trichopapilosus, which was
recorded from well-dated Valanginian
Hauterivian
sequences of northeast Libya (Thusu & van der Eem
1985; Thusu et al. 1988), occurs in most of these
samples.
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36
A.S. Deaf et al.
 Palynology 37
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Figure 6. The lithological succession in the Abu Tunis 1x borehole with sample positions, original age dating (WEPCO 1968),
key biostratigraphical events and revised ages as deduced in the current work.

J
Plate 1. Late Jurassic
Early Cretaceous spores and pollen grains. The sample/slide number and depth, England Finder coordi-
nates, and reference number of each taxon on the quantitative range chart (the latter in parentheses) are indicated for all speci-
mens; the scale bars represent 20 mm.
1. Dictyophyllidites harrisii Couper, 1958, slide AT-1A, 10,150 ft, O28/1, (25).
2. Dictyophyllidites sp., slide AT-3A, 10,050 ft, O20, (35).
3. Concavisporites sp., slide AT-6A, 9900 ft, Q32/2, (11).
4. Auritulinasporites scanicus Nilsson, 1958, silde AT-7A, 9850 ft, O11/2, (52).
5. Deltoidospora toralis (Leschik 1955) Lund, 1977, slide AT-1A, 10,150 ft, K34/3, (30).
6. Deltoidospora sp., slide AT-4A, 10,000 ft, S25, (1).
7. Deltoidospora hallii Miner, 1935, slide AT-3A, 10,050 ft, Q36/3, (5).
8. Triplanosporites sp., slide AT-3A, 10,050 ft, J42/1, (3).
9. Deltoidospora australis (Couper 1953) Pocock, 1970, slide AT-12A, 9600 ft, E33, (37).
10, 22. Taxacites sahariensis Reyre, 1973, slide AT-3B, 10,050 ft, O32, slide AT-10A, 9700 ft, P23/2, (71).
11. Balmeiopsis limbatus (Balme 1957) Archangelsky, 1979, slide AT-1A, 10,150 ft, L17/4, (60).
12. Araucariacites australis Cookson ex Couper, 1953, slide AT-1A, 10,150 ft, U16/2, (63).
13. Inaperturopollenites undulatus Weyland & Greifeld, 1953, slide AT-4A, 10,000 ft, M22/3, (65).
18. Classopollis sp. slide AT-10A, 9700 ft, M44, (64).
19. Classopollis classoides Pflug, 1953, slide AT-6A, 9900 ft, R30, (61).
23. Crybelosporites brenneri Playford, 1971, slide AT-3B, 10,050 ft, P32, (31).

Berriasian
Valanginian spores and pollen grains

17, 21. Pilosisporites trichopapillosus (Thiergrat 1949) Delcourt & Sprumont, 1955, slide AT-4A, 10,000 ft, M47/1, slide AT-5A,
9950 ft, M35, (55).
20. Impardecispora apiverrucata (Couper 1958) Venkatachala et al., 1969, slide AT-2A, 10,100 ft, M14, (59).

Late Hauterivian
Early Barremian spores and pollen grains

14, 15, 16. Dicheiropollis etruscus Trevisan 1972, slide AT-2B, 10,100 ft, J19/2, slide AT-3A, 10050 ft, M43/2, slide AT-3A, Q31/1, (76).
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38
A.S. Deaf et al.
 Palynology 39

In the palaeotropical region, Dicheiropollis etruscus
was recorded in northeast Libya from foraminifera-
and dinoflagellate-dated rocks of late Hauteri-
vian
early Barremian age (Thusu & van der Eem
1985; Thusu et al. 1988) and, similarly, from the paly-
nologically dated late Hauterivian
early Barremian
rocks of Senegal and the Ivory Coast (Salard-Chebol-
daeff 1990). In the palaeosubtropical region (§15
20
N and S), the earliest occurrence of D. etruscus is from
the base of turbiditic sediments of early Berriasian age
(G€ubeli et al. 1984), and its latest occurrence has been
recorded in Morocco from dinoflagellate-dated marine
sediments of early Barremian age (Hochuli 1981). In
Sudan, Gabon and other parts of the palaeosubtropics,
D. etruscus has been recorded from palynologically
dated continental and shallow marine sediments of
Neocomian
Barremian age (Jardine et al. 1974; Doyle
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The presence of Dicheiropollis etruscus, in forami-
nifera-dated late Hauterivian
early Barremian sedi-
ments in the palaeosubtropical African region,
provides strong evidence that samples 2
9 are of a late
Hauterivian
early Barremian age. Moreover, the
uppermost occurrence of D. etruscus in sample 9, just
below the first appearance of Stellatopollis spp., corre-
lates with the same event documented in the Dicheiro-
pollis/Afropollis Phytogeoprovince (Doyle et al. 1977;
G€ubeli et al. 1984; Thusu et al. 1988; Regali & Viana
1989; Figure 2), and supports an age not younger than
the early Barremian. As a result, a late Hauteri-
vian
early Barremian age is adopted for this interval
(Figure 6).
WEPCO (1968) referred to samples of this palyno-
zone as ‘no information’. The clastic (sandstone and
shales) lithology of samples 2 to 9 (10,100 to 9750 ft),

et al. 1977; Penny 1986; Kaska 1989; Salard-Chebol-
daeff 1990; Awad 1994). Similarly, D. etruscus has
been recorded in northeast Brazil from palynologically
dated fluvio-lacustrine rocks of Berriasian
early Bar-
remian age (Regali et al. 1974; Regali & Viana 1989).
The phytoplankton assemblage of Palynozone 1 is
represented by rare occurrences of some facies-
controlled ceratioid dinoflagellate species with pre--
Hauterivian and post-Barremian age ranges: Phobero-
cysta neocomica of late Berriasian
Barremian
stratigraphic range in the Tethyan Realm (Habib &
Drugg 1983; Thusu et al. 1988; Erba et al. 1999;
Torricelli 2000, 2001); Muderongia pariata, which has
an early Hauterivian
early Albian range in Italy (Tor-
ricelli 2000, 2001), and Pseudoceratium pelliferum,
which has a Tethyan late Berriasian
Barremian range
(Habib & Drugg 1983; Thusu et al. 1988; Hoede-
maeker & Leereveld 1995; Wilpshaar 1995; Leereveld
1997a; Torricelli 2000). Muderongia aequicorna is pres-
ent in the lower part of the interval, and is known from
the late Hauterivian in the Southern Alps of Italy (Tor-
ricelli 2000), where it was accurately dated by a variety
of means (Erba et al. 1999). This latter species repre-
sents the only marine evidence for a late Hauterivian
age proposed for the lower part of Palynozone 1.
with the above-assigned late Hauterivian
early Barre-
mian age, suggests that samples of Palynozone 1 repre-
sent the lower part of the Alam El Bueib Formation
(Figure 6). This formation is a clastic unit known to be
mainly composed of sandstone with frequent shale
interbeds in its lower part and occasional limestone
beds in its upper part. This unit is believed to range in
age at its type locality ‘the Alam El Bueib-1 well’
(Figure 1), from Berriasian to Aptian (Hantar 1990;
Ibrahim & Schrank 1996), and the environment of
deposition has been described as shallow marine, but
with a more continental influence toward the south
(Hantar 1990; Kerdany & Cherif 1990).
Correlation. The palynoflora of the current zone shows
a great similarity to the upper part of Miospore Associ-
ation V and the lower part of Miospore Association VI
(late Hauterivian
early Barremian) of Thusu et al.
(1988), northeast Libya. In the same regional context,
in the northern Western Desert of Egypt, the assem-
blage of ‘Section 1’ in the Mersa Matruh-1 well of
‘Neocomian’ and early Barremian age of Penny (1991),
and Zone II (late Hauterivian
early Barremian) of
Ibrahim & Schrank (1996) in the Kahraman-1 well, are
also similar to the present zone.

J
Plate 2. Late Barremian angiosperm pollen grains. The sample/slide number and depth, England Finder coordinates, and refer-
ence number of each taxon on the quantitative range chart (the latter in parentheses) are indicated for all specimens; the scale
bars represent 20 mm.
1, 2. Retimonocolpites ghazalii Ibrahim 2002, slide AT-16A, 9400 ft, U41/1, slide AT-24A, 9000 ft, K28/1, (91).
3, 4. Retimonocolpites matruhensis-Retimonocolpites ghazalii complex, slide AT-23B, 9050 ft, E28/3, slide AT-10A, 9700 ft, O21/4,
(93).
5, 8. Retimonocolpites matruhensis Penny 1986, slide AT-16A, 9400 ft, P26, slide AT-24A, 9000 ft, Q18/1, (92).
6, 10. Retimonocolpites bueibensis Ibrahim 2002, slide AT-6A, 9350 ft, J38, slide AT-18A, 9300 ft, V48/2, (96).
7. Retimonocolpites variplicatus Schrank & Mahmoud 1998, slide AT-14A, 9500 ft, P22, (101).
9, 12. Retimonocolpites pennyi Schrank & Mahmoud 2002, slide AT-19A, 9250 ft, W38, slide AT-16A, 9400 ft, O22, (97).
11, 13, 17. Dichastopollenites ghazalatensis Ibrahim 1996, slide AT-21B, 9150 ft, D15/4, slide AT-18B, 9300 ft, S37/3, slide AT-
20B, 9200 ft, M18/1, (98).
14, 19. Stellatopollis bituberensis Penny 1986, slide AT-19B, 9250 ft, E37/4, (99).
15, 16. Stellatopollis barghoornii Doyle in Doyle et al. 1975, slide AT-52B, 7600 ft, R30, slide AT-25B, 8950 ft, X29/2, (84).
18. Stellatopollis hughesii Penny 1986, slide AT-33B, 8550 ft, W37, (87).
 40 A.S. Deaf et al.
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Plate 3. Late Barremian angiosperm pollen grains. The sample/slide number and depth, England Finder coordinates, and refer-
ence number of each taxon on the quantitative range chart (the latter in parentheses) are indicated for all specimens; the scale
bars represent 20 mm.
1. Stellatopollis dejaxii Ibrahim 2002, slide AT-24A, 9000 ft, G13/2, (86).
2. Retimonocolpites sp.1 Schrank & Mahmoud 2002, slide AT-14A, 9500 ft, R30, (100).
3. Retiacolpites columellatus Schrank in Schrank & Mahmoud 2002, slide AT-21A, 9150 ft, Q23/1, (95).
4. Tucanopollis annulatus Schrank in Schrank & Mahmoud 2002, slide AT-17A, 9350 ft, K31/1, (94).
5-7. Afropollis operculatus Doyle et al. 1982, 5, slide AT-41A, 8150 ft, K22; 6, slide AT-49B,7750 ft, J31/3; 7, slide AT-40A, 8200
ft, Q48/4, (78).
15. Afropollis aff. zonatus Doyle et al. 1982, slide AT-20B, 9200 ft, R39, (80).

Early Aptian angiosperm pollen grains

8-10. Afropollis zonatus Doyle et al. 1982, slide AT-20B, 9200 ft, K35, slide AT-33A, 8550 ft, Y44, slide AT-23A, 9050 ft, Y35,
(85).

Late Aptian angiosperm pollen grains

11, 12. Afropollis aff. jardinus Doyle et al. 1982, slide AT-39A, 8250 ft, W29/3, slide AT-38A, 8300 ft, O24, (79).
13, 14. Afropollis sp. B Doyle et al. 1982, 1, slide AT-16B, 9400 ft, O23/3, slide AT-24B, 9000 ft, L31/4, (89).
 Palynology
5.2.3 Palynozone 2 (late Barremian):
Retimonocolpites spp.
Tucanopollis
annulatus
Pseudoceratium
retusum
Stellatopollis spp. Assemblage Zone
Samples. This zone is represented by samples 10 to 19
and covers a depth from 9700 to 9250 ft (2957
2819 m).
Definition. From the FAD of Retimonocolpites matru-
hensis (92), Retimonocolpites matruhensis-ghazalii com-
plex (93), Tucanopollis annulatus (94), Retimonocolpites
pennyi (97), Pseudoceratium retusum (123) and below
the FAD of Stellatopollis bituberensis (99) to just below
the FAD of Afropollis zonatus (85), Florentinia mantel-
lii (118) and Pseudoceratium securigerum (112).
Associated palynoflora. Balmeisporites longirimosus
(15), Microfoveolatosporites skottsbergii (17), Muro-
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spora cf. mesozoica (34), Leptolepidites major (42), Glei-
cheniidites rasilis (43), Leptolepidites psarosus (44),
Crybelosporites striatus (45), Aequitriradites verrucosus
(46), Echinatisporis varispinosus (53), Ephedripites spp.
(62), Reyrea polymorpha (67), Afropollis sp. B Doyle
et al. 1982 (89), Retimonocolpites variplicatus (101),
Subtilisphaera scabrata (126), Cribroperidinium edward-
sii (143), Circulodinium cf. attadalicum (151), Cyclone-
phelium cf. vannophorum (152) and Cyclonephelium
vannophorum (153).
Remarks. This interval shows a similarity in its palyno-
floral content with the underlying samples, but is also
characterised by a few forms of Ephedripites and thick-
walled spores, the incoming of true (columellate)
angiosperm pollen grains, e.g. Retimonocolpites
spp. and Stellatopollis spp. (Plates 2
3) and a few late
Barremian marker dinoflagellate species (Plate 4).
Age appraisal. The oldest records of Retimonocolpites
spp. and Stellatopollis spp. pollen are widely accepted
to mark the late Barremian in the Dicheiropollis/Afro-
pollis Phytogeoprovince (Penny 1991; Doyle 1992;
Penny 1992; Schrank 1992; Schrank & Mahmoud
1998; Doyle 1999). In the foraminifera- and dinoflagel-
late-dated Libyan late Barremian, Stellatopollis spp.
and Tucanopollis crisopolensis were recorded from the
earliest late Barremian, and Retimonocolpites spp. were
recorded from the latest late Barremian (Thusu & van
der Eem 1985; Thusu et al. 1988). In the well-dated
marine sediments of Morocco, Retimonocolpites spp.
and Stellatopollis spp. have been recorded from the
late Barremian (G€ ubeli et al. 1984). The heteropolar,
zonasulculate pollen grain Afropollis operculatus is
widely accepted as an Aptian marker species in the pre-
Albian Dicheiropollis etruscus/Afropollis Phytogeo-
graphical Province (Doyle et al. 1982; Uwins & Batten
41
1988; Salard-Cheboldaeff 1990; Schrank & Ibrahim
1995). Afropollis operculatus has been recorded (as
Reticulatasporites jardinus Type 1) from a planktonic
foraminifera- and calcareous nannoplankton-dated
early Aptian section at Deep Sea Drilling Project
(DSDP) Site 418B in the Western North Atlantic
(Hochuli 1981), from the partly ammonite-dated
Aptian Afropollis reference section of Gabon and the
benthic foraminifera- and ostracode-dated Aptian
Afropollis reference section of Senegal (Doyle et al.
1982; Doyle 1992), and from the supposed early Aptian
of Egypt (e.g. Schrank 1983; Schrank & Ibrahim 1995).
However, a pre-Aptian range of this species was
recorded in Morocco by G€ ubeli et al. (1984) from dino-
flagellate-dated marine rocks of late Barremian age,
and in the supposed late Barremian of Gabon, Egypt
and northeast Brazil (Regali & Viana 1989; Schrank &
Mahmoud 1998; Doyle 1999; Schrank & Mahmoud
2002). The biostratigraphical events indicating the late
Barremian of Egypt, Libya and Senegal are of the
same age as those of Morocco and Gabon. This simi-
larity in age of the bioevents implies the onset of a late
Barremian biostratigraphical isochroneity between
these palaeotropical and palaeosubtropical regions.
The base of this zone is characterised by a number
of angiosperm pollen grains first described from the
palynologically dated late Barremian of Egypt: Reti-
monocolpites matruhensis, Retimonocolpites pennyi,
Stellatopollis bituberensis, Tucanopollis annulatus,
Retiacolpites sp.1 and Retimonocolpites bueibensis
(Penny 1986; Penny 1991; Schrank & Mahmoud 2000;
Ibrahim 2002; Schrank & Mahmoud 2002). Stellato-
pollis barghoornii of probable late Barremian
Aptian
range in Egypt (Ibrahim & Schrank 1996; Ibrahim
2002) occurs in most of the samples from this interval.
In the upper part of Zone 2, Stellatopollis hughesii,
Stellatopollis dejaxii, Retimonocolpites ghazalii and
Afropollis aff. zonatus are all found, which were also
reported from the supposed late Barremian of Egypt
(Penny 1986; Ibrahim 2002). Afropollis aff. zonatus was
recognised in Gabon from the pre-salt early Aptian
rocks (Doyle et al. 1982; Doyle 1992) below the ammo-
nite-dated carbonate sequence of late Aptian
Albian
age (Reyment & Tait 1972), and from the palynologi-
cally dated early Aptian (Schrank 1983; Schrank &
Ibrahim 1995) and the late Barremian
early Aptian
(Penny 1986) of Egypt. The two possible late Barre-
mian pollen taxa Retiacolpites columellatus and Reti-
monocolpites sp.1 of Schrank & Mahmoud (2002) have
their FAD in the upper part of Palynozone 2.
However, in the middle part of this sample interval,
there are rare occurrences of Arecipites microfoveola-
tus, a taxon that was described by Ibrahim (2002)
from Aptian-age sediments from the Ghazalat-1
well. In addition, there are rare occurrences of
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42
A.S. Deaf et al.
 Palynology 43

Dichastopollenites ghazalatensis of Aptian
Cenoma-
nian age (Ibrahim 1996; Ibrahim 2002) in the upper
part of the interval, but until an independent extension
of the range of these two taxa into the late Barremian
is confirmed, their distribution in the present interval is
considered as extended due to possible caving.
In terms of dinoflagellate cysts, the lower boundary
of this zone is also marked by the FAD of the late
Barremian Tethyan species Pseudoceratium retusum
(Srivastava 1984; Wilpshaar 1995; Leereveld 1997b).
The phytoplankton assemblages also record the first
appearances of some Hauterivian
Barremian dinofla-
gellate species, including Subtilisphaera senegalensis
and Circulodinium brevispinosum, which have Tethyan
late Hauterivian
Albian ranges (Thusu et al. 1988;
Leereveld 1997b; Torricelli 2000, 2001). In the middle
part of the interval, two Tethyan latest Hauterivian/
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the late Barremian of Italy (Torricelli 2000) and ranges
into the early Albian. More problematic is the presence
of Florentinia cooksoniae, which was originally
described by Singh (1971) from the Albian lower
Shaftsbury Formation, Canada, and is also known
from the supposed Albian
Cenomanian rocks of
Egypt and northeast Libya (Uwins & Batten 1988;
Omran et al. 1990). However, the downward extension
of the range of F. cooksoniae into Palynozone 2 could
be a true stratigraphic occurrence, as this species was
recovered by Duxbury (1980) from the Barremian
Speeton Clay of England.
The appearance in this sample interval of dinofla-
gellate cyst taxa characteristic of the latest Hauteri-
vian/early Barremian
Aptian, together with several
late Barremian
early Aptian angiosperm pollen
grains, might suggest a late Barremian
early Aptian

early Barremian
Albian dinoflagellate species appear:
Subtilisphaera terrula (Habib & Drugg 1983; Torricelli
2001) and Subtilisphaera perlucida (Habib & Drugg
1983; Thusu et al. 1988; Leereveld 1997b; Torricelli
2000, 2001). Subtilisphaera scabrata, used by Leereveld
(1997b) to delineate the lower boundary of his Tethyan
Ssc Zone of early Barremian age, also occurs in the
middle part of the interval. The upper part of the
interval is characterised by the appearance of Pseudo-
ceratium anaphrissum, regarded as a late Barre-
mian
Aptian marker species in the Tethyan Realm
(Thusu et al. 1988; Foucher et al. 1994; Hoedemaeker
& Leereveld 1995; Figure 2). The inception of Odonto-
chitina operculata was used in the Tethyan Realm to
determine the base of the late Barremian (Wilpshaar
1995; Leereveld 1997b; Torricelli 2000, 2001), and
appears in the uppermost part of this sample interval,
as does Odontochitina ancala, which was recorded from
age. However, in the overlying sample (20), the pres-
ence of the Aptian angiosperm pollen marker Afropol-
lis zonatus and the characteristic early Aptian
dinoflagellates Pseudoceratium securigerum and Palae-
operidinium cretaceum indicates that this interval
should be dated as late Barremian. The mixed, clastic-
carbonate lithology of samples 10
19 (9700
9250 ft)
and their age identified here suggest that samples of
Palynozone 2 comprise the upper part of the Alam El
Bueib Formation.
Correlation. The angiosperm pollen assemblage (Stella-
topollis
Retimonocolpites
Afropollis) recorded in this
zone is identical to that described in Zones CVI
CVII
(late Barremian) of Doyle et al. (1977), northeast
Gabon, Zone D (late Barremian) of G€ ubeli et al.
(1984), northern Morocco, Miospore Association VI
(late Barremian) of Thusu et al. (1988), northeast Libya,

J
Plate 4. Late Barremian dinoflagellate cysts. The sample/slide number and depth, England Finder coordinates, and reference
number of each taxon on the quantitative range chart (the latter in parentheses) are indicated for all specimens; the scale bars rep-
resent 20 mm.
1. Pseudoceratium anaphrissum (Sarjeant 1966) Bint, 1986, slide AT-27A, 8850 ft, U36/2, (119).
2,10. Pseudoceratium retusum Brideaux, 1977, slide AT-24A, 9000 ft, J39/4, slide AT-21A, 9150 ft, R38/3, (123).
7. Odontochitina ancala Bint, 1986, slide AT-29A, 8750 ft, X29, (138).
8. Odontochitina operculata (Wetzel 1933) Deflandre & Cookson, 1955, AT-19B, 9250 ft, U30/2, (121).

Early Cretaceous dinoflagellate cysts

3, 5. Subtilisphaera senegalensis Jain & Millepied, 1973, slide AT-22B, 9100 ft, W36, slide AT-28A, 8800 ft, K20/2, (113).
4, 9. Subtilisphaera perlucida (Alberti 1959) Jain & Millepied, 1973, slide AT-22A, 9100 ft, O32, slide AT-28A, 8800 ft, S29 (117).
6. Subtilisphaera terrula (Davey 1974) Lentin & Williams, 1976, slide AT-31A, 8650 ft, W37, (130).

Cretaceous dinoflagellate cysts

11. Downiesphaeridium sp., slide AT-34A, 8500 ft, U14/1, (131).
15. Coronifera albertii Millioud, 1969, slide AT-24A, 9000 ft, T35/2, (147).
16. Oligosphaeridium complex (White 1842) Davey & Williams, 1966, slide AT-25A, 8950 ft, L43, (110).
17. Oligosphaeridium albertense (Pocock 1962) Davey & Williams, 1969, slide AT-29A, 8750 ft, Q47, (134).
18. Oligosphaeridium poculum Jain, 1977, slide AT-23B, 9050 ft, N22/2, (115).

Acritarchs
12. Veryhachium collectum Wall, 1965, slide AT-33A, 8550 ft, X39/4, (160).
13. Veryhachium metum Davey, 1970, slide AT-42A, 8100 ft, T31/3, (161).
14. Micrhystridium stellatum Deflandre, 1945, slide AT-37A, 8350 ft, O40/4, (162).
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44
A.S. Deaf et al.
 Palynology 45

and the Stellatopollis bituberensis Zone (late Barremian)
of Regali & Viana (1989), northeast Brazil. In the north-
ern Western Desert of Egypt, the assemblage of Section
2 (late Barremian) of Penny (1991) in the Mersa
Matruh-1 well, the lower part of Zone IV (late Barre-
mian
early Aptian) of Ibrahim & Schrank (1996) in
the Kahraman-1 well, and assemblage ‘A1’ of late
Barremian age of Ibrahim (2002) from the Ghazalat-1
well, also bear a striking similarity to the current zone.
5.2.4 Palynozone 3 (Aptian) Afropollis
zonatus
Afropollis operculatus

Pseudoceratium securigerum

Palaeoperidinium cretaceum Assemblage Zone
Samples. This zone includes samples from 20 to 57,
which were taken from depths between 9200 and 7350
ft (2804
2240 m).
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phytoplankton increases in abundance and diversity
from a maximum of 155 (avg. 33) cysts/gram compris-
ing some 16 species in the underlying interval, to a
maximum 1171 (avg. 100) cysts/gram, and with a diver-
sity of some 25 species in this sample interval (Figure 5).
However, pteridophyte spores, gymnosperms and the
angiosperm pollen found in the underlying Barremian
interval still dominate the palynofloral assemblage.
Age appraisal. The base of this interval is characterised
by the incoming of the isopolar zonasulculate pollen
Afropollis zonatus (Plate 3, figures 8
10), which is
regarded as an early Aptian marker species in the
Dicheiropollis etruscus/Afropollis Phytogeographical
Province (Figure 2). A. zonatus was first described
from palynologically dated sediments of early Aptian
age in Gabon (Doyle et al. 1982; Doyle 1992), and was

later recorded (as Reticulatasporites jardinus Type 2)

Definition. From the FAD of Afropollis zonatus (85),
Pseudoceratium securigerum (112) and Florentinia man-
tellii (118) and just below the FAD of Palaeoperidinium
cretaceum (114) to just below the FAD of Afropollis
jardinus.
Associated palynoflora. Crybelosporites pannuceus (2),
Cicatricosisporites orbiculatus (4), Cicatricosisporites
sinuosus (6), Murospora florida (7), Triporoletes reticu-
latus (9), Balmeisporites cf. holodictyus (13), Microfo-
veolatosporites skottsbergii (17), Matonisporites spp.
(19), Kyrtomisporis spp. (24), Aequitriradites norrissii
(39), Gemmatrilites sp. (40), Januasporites sp. (41), Tri-
colpites sp. (81), Tricolpites vulgaris (90), Oligosphaeri-
dium complex (110), Florentinia spp. (111),
Oligosphaeridium poculum (115), Coronifera tubulosa
(128), Downiesphaeridium spp. (131), Coronifera alber-
tii (147), Pseudoceratium almohadense (149).
Remarks. The samples of this interval differ in their
floral content from the underlying intervals, as
from foraminiferally dated rocks of the same age in
southern Switzerland (Hochuli 1981) and from dinofla-
gellate-dated sediments in northern Morocco (G€ ubeli
et al. 1984).
Afropollis aff. jardinus (Plate 3, figures 11
12),
which was reported from palynologically dated rocks
of late Aptian
early Albian age in West Africa
and Egypt (Doyle et al. 1982; Penny 1989; Schrank &
Ibrahim 1995; Ibrahim & Schrank 1996), also occurs in
this interval.
However, overall, this interval contains relatively
few angiosperm pollen grains that could be viewed as
Aptian markers, and it is the dinoflagellates that are of
greater biostratigraphical importance. The base of this
interval is also delineated by the FAD of Pseudocera-
tium securigerum, a diagnostic Tethyan early Aptian
form (Figure 2; Plate 5, figures 1
2), which has been
recorded from the base of the ammonite-dated Aptian
type section in southeast France (Davey & Verdier
1974), from the foraminifer-dated Aptian of Algeria
(Foucher et al. 1994) and from the dinoflagellate-dated

J
Plate 5. Early Aptian dinoflagellate cysts. The sample/slide number and depth, England Finder coordinates, and reference num-
ber of each taxon on the quantitative range chart (the latter in parentheses) are indicated for all specimens; the scale bars repre-
sent 20 mm.
1, 2. Pseudoceratium securigerum (Davey & Verdier 1974) Bint, 1986, slide AT-20A, 9200 ft, T40/3, slide AT-20B, Q34, (112).
4, 9. Palaeoperidinium cretaceum (Pocock 1962) Lentin & Williams, 1976, slide AT-36A, 8400 ft, R12, slide AT-21A, 9150 ft,
M41, (114).
5, 6, 14. Florentinia mantellii (Davey & Williams 1966) Davey & Verdier 1973, slide AT-22A, 9100 ft, Q51, slide AT-32A, 8600 ft,
W36, slide AT-19A, 9250 ft, S23/1, (118).
13. Florentinia laciniata Davey & Verdier 1973, slide AT-23A, 9050 ft, N37, (145).
17. Aptea polymorpha Eisenack, 1958, slide AT-24A, 9000 ft, N30/3, (129).

Early Cretaceous dinoflagellate cysts

3, 7. Subtilisphaera scabrata Jain & Millepied, 1973, slide AT-20A, 9200 ft, M45/1, slide AT-29A, 8750 ft, U46, (126).
8, 11. Spiniferites sp., slide AT-24A, 9000 ft, Q27/2, slide AT-23A, 9050 ft, S43/3, (116).
10. Cribroperidinium edwardsii (Cookson & Eisenack 1958) Davey, 1969, slide AT-27, 8850 ft, H42/3, (143).
12. Florentinia cooksoniae (Singh 1971) Duxbury 1980, slide AT-21A, 9150 ft, K30/3, (139).
15. Cyclonephelium vannophorum Davey, 1969, slide AT-12A, 9600 ft, K28/4, (153).
16. Pseudoceratium almohadense (Below 1984) Lentin & Williams, 1989, slide AT-23A, 9050 ft, X42/2, (149).
 46 A.S. Deaf et al.
marine sediments of southern Italy (Torricelli 2001). In
southern England, P. securigerum was also recovered
by Duxbury (1983) and Lister & Batten (1988) from
ammonite-calibrated boreal marine sediments of early
Aptian age. In northwest Egypt and northeast Libya,
this same species was recovered from rocks of pre-
sumed early Aptian age (Uwins & Batten 1988; Omran
et al. 1990; El-Beialy 1994; Schrank & Ibrahim 1995).
Another early Aptian marker form with its FAD
just above the base of this sample interval is Palaeoper-
idinium cretaceum. This species has been recovered
from rocks that were age calibrated by ammonites,
planktonic foraminifera and calcareous nannoplank-
ton as early Aptian in the Tethyan Realm of southeast
France and Italy (Davey & Verdier 1974; Torricelli
2000, 2001 2006); from dinoflagellate-dated samples
from DSDP Site 543A in the Western Central Atlantic
Downloaded by [Assiut University] at 08:01 24 February 2016
(Habib & Drugg 1983) and from the supposed early
Aptian of Egypt and northeast Libya (Uwins & Batten
1988; Ibrahim et al. 2002). The later appearance of
Aptea polymorpha in sample 24 above the lower part of
this sample interval is consistent with its FAD in the
late early Aptian in the Tethyan Realm (Figure 3) and
elsewhere (Davey & Verdier 1974; Lister & Batten
1988; Foucher et al. 1994; Torricelli 2000, 2001). Other
characteristic Aptian dinoflagellate species which
appear in this interval are: Florentinia laciniata,
recorded from ammonite-dated rocks of Aptian
early
Cenomanian age in southeast France (Davey & Verdier
1973), and from well-dated Aptian rocks of Italy (Tor-
ricelli 2001); and Florentinia mantellii, also of Aptian
aspect, being recorded from the ammonite-dated late
Aptian of southeast France (Davey & Verdier 1974),
and the dinoflagellate-dated late Aptian of Italy (Torri-
celli 2000, 2001).
Based on the Aptian dinoflagellate and angiosperm
markers, and with the FAD of the diagnostic
early Albian angiosperm Afropollis jardinus in the over-
lying sample 58, an Aptian age is suggested for this
interval.
The mainly dolomitic unit represented by samples
20
41 (9200
8150 ft), and the assigned Aptian age
herein, suggest this part of the studied sequence is the
Alamein Formation. This formation is widely distrib-
uted over northern Egypt, and is composed at its type
locality ‘the Alamein-1 well’ (Figure 1) of a light
brown, hard microcrystalline dolomite with vuggy
porosity. The Alamein Formation has been attributed
an Aptian age, and seems to have been deposited in a
shallow marine, low- to moderate-energy environment
(Kerdany & Cherif 1990). However, the upper clastic
part of this palynozone (samples 42
57, 8100
7350 ft)
represents the Dahab Formation. This formation is
known from its type locality ‘the Dahab-1 well’
(Figure 1) in the north Western Desert where it is
composed of a shale unit interbedded with thin part-
ings of siltstone and sandstone. The Dahab Formation
is attributed an Aptian
early Albian age, and it seems
to have been deposited in a shallow marine environ-
ment (Barakat & Darwish 1982; Hantar 1990).
Correlation. The majority of the angiosperm pollen
and dinoflagellate cyst index taxa of this zone corre-
spond closely to that described in Assemblage II
(early
late mid Aptian) of Uwins & Batten (1988),
northeast Libya and the Aptian Zone E/lower part of
Zone F of G€ ubeli et al. (1984), northern Morocco.
Zones CVII
CIX (Aptian) of Doyle et al. (1977) from
northeast Gabon and the Exesipollenites tumulus Zone
(Aptian) of Regali & Viana (1989) from northeast
Brazil also share angiosperm pollen index taxa with
Palynozone 3. In the northern Western Desert of
Egypt, some angiosperm-based zonal schemes corre-
spond to this zone: e.g. the Aptian Afropollis opercula-
tus
Brenneripollis
Tricolpites spp. Assemblage Zone
of Schrank & Ibrahim (1995) in the Kahraman-1 well,
and assemblages ‘A2’ and ‘A3’ of early
mid Aptian
age of Ibrahim (2002) in the Ghazalat-1 well (which
only equates with the lower part of Palynozone 3). The
Aptian pollen-spore (PSIII) and dinoflagellate cysts
(D2) zones of Mahmoud & Deaf (2007) identified in
the Siqeifa 1-X borehole are the only ones that closely
resemble the current zone in their composition of both
terrestrial and marine palynomorphs.
6. Discussion
Due to the close palynofloral and palaeogeographic
relationships, this section will focus on specific compar-
isons between the palynostratigraphy of the Abu Tunis
1x well with results from West Africa (Senegal, the
Ivory Coast and Nigeria) and North Africa (northwest
Egypt, northeast Libya and northern Morocco).
6.1. Quantitative pre-Albian palynofloral characteris-
tics in the Abu Tunis 1x well
The palynofloras recorded from the late Hauteri-
vian
Aptian interval in the Abu Tunis 1x well reveal
all of the diagnostic characteristics of the pre-Albian
Dicheiropollis/Afropollis Phytogeographical Province
as recorded from other locations in equatorial Africa.
These include high abundances (4
90%, avg. 24%;
1045
32,475 grains/g, avg. 7480) and a moderate
diversity (12 species) of smooth trilete spores (e.g. Del-
toidospora and Concavisporites), the appearance of
Dicheiropollis in low occurrences (0.4
1.2%, avg.
0.8%; 23
110, avg. 65 grains/g), the diversity of Ephe-
dripites (12 species) in the late Aptian and an absence
of bi- and tri-saccate pollen grains. Abu Tunis 1x also
yields low occurrences of Taxacites (D Exesipollenites)
 Palynology 47

sahariensis (1
13%, avg. 4%; 21
293, avg. 95 grains/g)

and moderate abundances of Araucariacites (2
20%,
avg. 12%; 28
796, avg. 367 grains/g), in addition to
the presence of Tucanopollis (1
2%, avg. 1%; 11
33,
avg. 18 grains/g) and the appearance and diversity (five
species) of Afropollis (2
14%, avg. 5% of all total paly-
nomorphs; 78
2584, avg. 640 grains/g).

6.2. The palaeoecology and stratigraphic range of

Dicheiropollis etruscus
Trevisan (1972) noted that Dicheiropollis etruscus was
of cheirolepidiacean conifer affinity, and, as with
Classopollis, may have been both thermophilous and
adapted to arid conditions. High abundances of Class-
opollis are associated with evaporites, salts and redbed
deposits, and also with xeromorphic cheirolepidiacean
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wood and leaf megafossils, which further support hot,

dry conditions for this genus (Watson 1988; Doyle
1999). Classopollis was most abundant during Barre-
mian
Aptian time in the hot, dry subtropical latitudes
(15
30 N and S of the palaeoequator), while it is
found in lower occurrences in the hot, but slightly wet-
ter, tropical region (Doyle 1999). Doyle et al. (1982),
Schrank (1990) and Brenner (1996) all suggested rela-
tively wetter palaeoclimates for the African tropics
(e.g. Egypt and Sudan), based on the presence of high
abundances of fern spores (indicating humidity), and
lower frequencies of Classopollis and the cooler-tem-
perature coniferous genus Araucariacites than seen in
the subtropics.
In the subtropical region, several authors have
reported an earlier (Berriasian) appearance of Dicheir-
opollis etruscus, for example in the palynologically
dated continental clastic and salt deposits of West
Africa (Gabon: Doyle et al. 1977), from the Berriasian
of northern Morocco (G€ ubeli et al. 1984) and in the
subtropical region of northern South America (Muller
et al. 1987; Regali 1989). These records contrast
markedly with the species’ delayed late Hauterivian
appearance in the tropical northeast African region
(e.g. Thusu & van der Eem 1985; Uwins & Batten
1988, Libya; Figure 7). As Dicheiropollis etruscus is
known to have a late Hauterivian
early Barremian
range in north East Africa in foraminifer- and dinofla-
gellate-dated sediments (Libya: Uwins & Batten 1988),
this latter range is therefore accepted here as applicable
to Egypt and thus used to allocate a late Hauteri-
vian
early Barremian age for Palynozone 1 of the
Abu Tunis 1x borehole. The difference in the first
appearance data in these two regions could be inter-
preted in one of two ways: Figure 7. Early Cretaceous palynological correlation of
North and West Africa and northern South America (modi-
fied from Schrank 1992) and the palynological zonation pro-
(1) That the northeast and West African sequences
posed in the present study.
are diachronous
but this suggestion cannot
 48 A.S. Deaf et al.
be proven until independent age controls are
provided for D. etruscus in West Africa or
northern South America.
(2) The taxon’s distribution is palaeoecologically
controlled in each area. Prior to the breakup of
Western Gondwana; Egypt, Libya, Senegal
and the Ivory Coast lay approximately at the
palaeoequator, where as Morocco and Gabon
were located at subtropical latitudes around
10
15 N and S, respectively. D. etruscus thus
first appeared in hot, dry subtropical African
regions, but as Western Gondwana broke up
and the African Plate moved northeast during
the Late Jurassic and onward, Egypt, Libya
and Sudan were, by the late Hauterivian,
moved into a palaeo-subtropical position
(Figures 3
4). Hence, the more arid conditions
Downloaded by [Assiut University] at 08:01 24 February 2016
probably allowed D. etruscus to migrate into
these areas.
6.3. Aptian Afropollis acme events and local palaeocli-
matical effects
Although widely separated palaeogeographic areas,
Senegal and Egypt were located at more or less the
same palaeolatitude during the Aptian, and demon-
strate similar angiosperm-dominated assemblages of
very similar biostratigraphical ranges. Egypt and Sene-
gal were bordered by the Tethyan and Southern Atlan-
tic Oceans, respectively. As oceans are known as the
main driver for global (palaeo-) climate, they would be
expected to have similar ecological effects on the abun-
dance of some of the angiosperm-producing plants.
Doyle et al. (1982) recorded a first acme of Afropol-
lis (2
7% of total palynomorphs) in the foraminifera-
dated Aptian of Senegal, where persistent occurrences
of A. operculatus and A. zonatus were reported to range
from the early to late Aptian. This bears a striking sim-
ilarity to the Aptian acme of Afropollis (2
14%, avg.
5% of all total palynomorphs; 78
2584, avg. 640
grains/g) recorded in this study, also manifested by per-
sistent occurrences of A. zonatus and A. operculatus
(with even higher frequencies of the latter) into the late
Aptian (Figure 8). A similar first acme of Afropollis
(4
37%, avg. 9.5% of total palynomorphs) in the sup-
posed Aptian of Egypt has been also reported by
Schrank & Ibrahim (1995), but with no quantitative
species differentiation. This further supports the view
that the Aptian sequences of Senegal and Egypt are
indeed synchronous. Doyle et al. (1982) attributed the
upward persistence of A. operculatus and A. zonatus to
palaeoclimatical conditions, suggesting that Afropollis-
producing plants were better adapted to wetter, proba-
bly humid, coastal environments. In contrast to this, in
Gabon, there is an upward decline in Afropollis from
the pre-salt early Aptian Zone C-VII into the late
Aptian salt sequence Zones C-VIII
C-IX. This change
was explained by Doyle et al. (1982) as due to increased
aridity.
The wet coastal conditions proposed for the Afro-
pollis parent plants were later supported by Schrank
(2001), where he reported exceptionally high relative
abundances of Afropollis (35
78% of total palyno-
morphs) and elaterate pollen (11
15% of total palyno-
morphs) from Albian
Cenomanian continental
sediments of northern Sudan, which also yielded dino-
flagellate cysts believed to be tolerant of low salinities.
Schrank (2001) compared these extraordinary abun-
dances of Afropollis and elaterates to a similar event
recorded by El-Shamma (1991) from the marine
Albian
Cenomanian sediments of northern Egypt,
and suggested that the parent plants of both Afropollis
and elaterate pollen may have flourished in humid
coastal habitats. The temporary humid conditions
would have been brought to the intracontinental basins
of Sudan by a short-lived transgression.
Such observations may permit the recognition that
high abundances of Afropollis can be used as a proxy
indicator for warm, humid coastal conditions.
6.4. A unified Early Cretaceous palynological zonal
scheme for the northern Western Desert of Egypt
Comparison between the zonal scheme erected here
with those proposed by Schrank & Ibrahim (1995) and
Ibrahim & Schrank (1996) for the Early Cretaceous of
the northern Western Desert shows great consistency
in zonal concept and composition for the late Hauteri-
vian
early Barremian and Aptian time intervals
(Figure 7), thus allowing a unification of these
biozones.
The Late Jurassic
Early Cretaceous hiatus, which
occurred in some basins of the northern Western Des-
ert (including the well studied here), prevents the erec-
tion of a complete biozonation for pre-late
Hauterivian sequences in this part of Egyptian stratig-
raphy, so there is a need for further studies of this time
interval preserved in other areas. However, the late
Barremian of Egypt is characterised by several angio-
sperm diagnostic taxa, notably Retimonocolpites and
Stellatopollis, that have been widely encountered on a
local scale (see Section 5.2.3). Species of these genera
would comprise at least a potentially important Egyp-
tian biozone for the late Barremian. Certain species of
these genera have also been recorded at regional (e.g.
Libya: Thusu et al. 1988) and interregional (e.g.
Gabon: Doyle et al. 1977) scales. However, much work
is still needed to document and calibrate the rest of the
Retimonocolpites and Stellatopollis-belonging species
before considering a local informal zonation.
 Palynology 49
Downloaded by [Assiut University] at 08:01 24 February 2016

Figure 8. Quantitative distribution of Afropollis and its acme events in Egypt and Senegal.
 50 A.S. Deaf et al.
7. Conclusions
This paper provides the first unified quantitative and
semi-quantitative pre-Albian palynofloral abundances
for an Egyptian succession, which permits the wider
regional and interregional correlations of palynofloral
assemblages. The sporomorphs recovered from the
Abu Tunis 1x borehole exhibit the characters of the
pre-Albian Dicheiropollis/Afropollis Phytogeographical
Provinces of equatorial Africa. The vertical distribu-
tion of index palynomorph taxa in the Abu Tunis 1x
has enabled the recognition of three palynological bio-
zones corresponding to three Early Cretaceous lithos-
tratigraphical units that have proved difficult to date
accurately in the past.
Correlation of the early Cretaceous zonal scheme
proposed for the Abu Tunis 1x well with those pro-
posed by Schrank & Ibrahim (1995) and Ibrahim &
Downloaded by [Assiut University] at 08:01 24 February 2016
Schrank (1996) for the northern Western Desert has
proved that the late Hauterivian
early Barremian and
Aptian share a common biostratigraphy. However, it
is still not possible to propose a complete Egyptian
pre-late Hauterivian biozonation due to hiatuses in
many of the studied successions. Further work is still
required to better calibrate the stratigraphical ranges
of the late Barremian species.
Correlation of the Early Cretaceous palynomorph
assemblages and biostratigraphical events in the Abu
Tunis 1x borehole with their palaeotropical, contempo-
raneous regional (Libya) and interregional (Senegal
and the Ivory Coast) equivalents has revealed a broad
similarity in the biostratigraphical range (late Hauteri-
vian
early Barremian) of the pollen index grain
Dicheiropollis etruscus in these areas.
The questionable Berriasian biostratigraphical
appearance of Dicheiropollis etruscus recorded from
West and northwest African palaeosubtropical coun-
tries (e.g. Gabon and Morocco) is suggested here to be
less likely due to a diachroneity between the sedimen-
tary sequences of both northeast and West Africa, as
there is as yet no (bio)- or litho-stratigraphic evidence
to support this. We suggest that this discrepancy in the
biostratigraphical appearances of D. etruscus could be
due to ecological factors driven by palaeolatitudinal
positions (i.e. the dry palaeosubtropical of West Afri-
can countries versus the relatively wetter palaeotropi-
cal countries of northeast Africa), where a dry
palaeoclimate is suggested as controlling this earlier
appearance. Hence, D. etruscus appeared earlier (Ber-
riasian) in northwest Africa and northern South Amer-
ica but migrated into northeast African regions in the
Hauterivian due to the northward movement of the
African Plate towards Laurasia.
The uppermost Barremian
Aptian successions in
Egypt and Senegal were deposited isochronously, and
this is supported by the recognition of coeval acme
events of Afropollis in both regions. The persistent
abundances of Afropollis zonatus and Afropollis oper-
culatus into the late Aptian of the Abu Tunis 1x bore-
hole (Palynozone 3) are believed to be due to a more
favourable, wetter palaeoclimate, and it is suggested
that high abundances of Afropollis may be used as a
proxy indicator for warm, humid coastal conditions.
Acknowledgements
The authors wish to thank the Egyptian General Petroleum
Corporation for providing well logs and samples from the
Abu Tunis 1x borehole. Mr. Shir Akbari at National Ocean-
ography Centre Southampton (NOCS), University of South-
ampton, is also thanked for his help in the laboratory.
Supplemental data
Supplemental data for this article can be accessed here http://
10.1080/01916122.2014.993480.
Author biographies
AMR S. DEAF was appointed as a lec-
turer in Palynology, Stratigraphy and
Palaeoecology at the Department of
Geology, Assiut University, Egypt in
2010. He received a BSc in geology in
1997, and an MSc in Palynology during
2003, from the same institution. Amr
obtained his PhD for research on the
palynostratigraphy, palaeoclimate,
palaeoecology and hydrocarbon potential of the Cretaceous
rocks of northern Egypt, from the National Oceanography
Centre, University of Southampton, U.K. in 2010. His
research interests are palynology, stratigraphy, palaeoecol-
ogy, palaeoclimatology, palaeoceanography and petroleum
geology, specially on the Mesozoic of the Middle East. Qua-
ternary geology of Egypt and other West African countries is
another area of research interest.
IAN C. HARDING is a reader in Palae-
oceanography, Micropalaeontology, and
Palynology at the National Oceanogra-
phy Centre, University of Southampton,
U.K. He received his BSc in Geology
from Nottingham University in 1982,
and his PhD in Micropalaeontology
from Cambridge University in 1986. His
research interest is focused on palaeocli-
mate change and palaeoenvironmental reconstruction using
micropalaeontological (palynological) proxies, especially in the
Paleogene. The development and integration of dinoflagellate
cysts, palynofacies analysis and geochemical indices as palaeo-
ceanographic proxies is a central research goal, and he has spe-
cial interest in hyperthermal events and greenhouse-icehouse
transition, and the sedimentological and micropalaeontological
investigation of taphonomic processes involved in exceptional
fossil preservation, especially high-resolution analysis of lami-
nated and finely bedded sediments. Ian was the Chair of Paly-
nology Group of The Micropalaeontological Society (TMS)
 Palynology
from 2005 to 2010, and the TMS representative on the board
of the International Federation of Palynological Societies from
2008 to 2010. He was Director at Large of the American Asso-
ciation of Stratigraphic Palynologists-The Palynology Society
from 2009 to 2011, President 2012
2013, and Past President
2013
2014.
JOHN E.A. MARSHALL is a Profes-
sor of Earth Science at the National
Oceanography Centre, University of
Southampton, U.K. John obtained a
BA in Natural Sciences from University
of Cambridge in 1976, and his PhD
from the Department of Botany, Uni-
versity of Bristol in 1981. He subse-
quently worked at the University of
Newcastle upon Tyne and Gearhart Geodata in Aberdeen.
His research interests are Mid Palaeozoic mass extinction
events, Devonian terrestrial palaeoclimates and the Earth
System, the spread of the Devonian forests, Devonian oil and
Downloaded by [Assiut University] at 08:01 24 February 2016
gas reservoirs, early seed-plants, the age and environment of
the early tetrapods, organic matter-waste water interactions
in waste tips and thermal maturity determination including
vitrinite and chitinozoan reflectivity and quantitative spore
colour. He has worked particularly in South America, Green-
land, Scotland, Spitsbergen and China. John was the Presi-
dent of the Commission Internationale de la Microflore du
Paleozo€ıque (CIMP) from 2006 to 2010, and a visiting Pro-
fessor at the Chinese Academy of Sciences, Nanjing, China
from 2010 to 2011. He is currently the Chair of the Sub-Com-
mission on Devonian Stratigraphy and Palynology Subject
Editor of the Journal of Micropalaeontology.
ORCID
Amr S. Deaf http://orcid.org/0000-0002-5073-7911
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