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Author(s): Amr S. Deaf, Ian C. Harding and John E. A. Marshall

Article title: Cretaceous (Albian–early? Santonian) palynology and stratigraphy of the Abu Tunis 1xborehole, northern Western Desert, Egypt

Article no: 828662

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Cretaceous (Albian–early? Santonian) palynology and stratigraphy of the Abu Tunis 1x borehole,northern Western Desert, Egypt

Amr S. Deafa,b*, Ian C. Hardinga and John E. A. Marshalla

aSchool of Ocean & Earth Science, National Oceanography Centre, Southampton (NOCS), University of Southampton, EuropeanWay, Southampton, SO14 3ZH, UK; bGeology Department, Faculty of Science, Assiut University, Assiut, 71516, Egypt

A palynological analysis has been conducted on the middle-upper Cretaceous sediments of the Abu Tunis 1xborehole, in the northern Western Desert, Egypt. The recovered palynomorphs have been analysed both qualita-tively and quantitatively and permit a refinement of the original stratigraphy with the identification of four time-rock units – these have been divided into four informal sporomorph and one dinoflagellate cyst palynozones. Thesebiozones are, from oldest to youngest: the Afropollis jardinus-Tricolporopollenites-Elaterosporites klaszii AssemblageZone (early–mid Albian), the Elaterosporites verrucatus-Sofrepites legouxae-Cretacaeiporites Assemblage Zone (lateAlbian–early Cenomanian), the Sofrepites legouxae Partial Range Zone (early–?mid Cenomanian), the Proteaciditescf. africaensis Total Range Zone (mid–late Cenomanian), and the Canningia senonica Total Range Zone (early?Santonian). A barren interzone has been identified just below the youngest palynozone, and this may be related tothe unfavourable lithology (i.e., limestone and dolostone).

The absolute abundance spore and pollen data represents the first quantitative description of an EgyptianAlbian–Cenomanian palynoflora, a flora that is characteristic of the Albian–Cenomanian Elaterate PhytogeographicProvince. The early Santonian palynoflora is exclusively marine phytoplankton; terrestrial palynomorphsrepresentative of the Senonian Palmae Province are completely absent. The quantitative and semi-quantitativedistributions of Afropollis jardinus have been compared with similar semi-quantitative distributions of this speciesfrom other wells in the north Western Desert of Egypt, and this permitted the identification of a mid Albian–earlyCenomanian Afropollis jardinus ‘acme’ as an important local biostratigraphic event in the mid Cretaceous.

Keywords: Cretaceous; palynology; stratigraphy; Western Desert; Egypt

1. Introduction

Egypt is currently located in the subtropical arid zone

and thus all its exposed sediments have been subjected

to extensive and deep weathering and are therefore paly-nologically barren. Most of the palynological research

conducted on Egyptian Cretaceous successions has been

based on deep borehole samples taken from exploratory

boreholes (e.g., Abdelmalik et al. 1981; Penny 1986;

Omran et al. 1990; Schrank & Ibrahim 1995; Mahmoud

& Deaf 2007; El Soughier et al. 2010), which were

drilled during hydrocarbon exploration activity in the

northern part of the EgyptianWestern Desert.The middle Cretaceous (Albian–Cenomanian) mixed

clastic-carbonate rocks represented by the Kharita,

Bahariya and lower Abu Roash formations of the north-

ern basinal area of Egypt were mainly deposited in very

shallow marine (brackish to coastal) to inner neritic open

marine conditions (Hantar 1990; Kerdany & Cherif

1990). These conditions were unfavourable for the prolif-

eration of planktonic forams and calcareous nannofos-sils and, thus, no independent age control is available, at

least for the Kharita succession. The upper Cretaceous

interval is generally composed of pre-Campanian (Turo-

nian–Santonian) middle to upper shelf deposits (upper

Abu Roash and Khoman ‘B’ formations), and of deeper

upper to middle slope deposits (Khoman ‘A’ Formation)

for the Campanian–Maastrichtian interval, which is

mainly represented by thick carbonate successions(Hantar 1990; Kerdany & Cherif 1990). Most of the

independently calibrated palynological work has been

produced as unpublished internal reports by oil explora-

tion companies. Thus the planktonic foraminifera-cali-

brated palynological work of Schrank and Ibrahim

(1995) and that of Abdel-Kireem et al. (1996) on upper

Cretaceous (Cenomanian–Maastrichtian) samples from

the Kahraman-1 and Abu Gharadig-1 boreholes in thenorthern Western Desert of Egypt is one of the few

attempts to provide micropalaeontologically calibrated

palynological work. Our own attempts to provide an

independent calibration using calcareous nannofossils

has proven unsuccessful: smear slides made from 10 sam-

ples of the carbonate succession of the Abu Tunis 1x

borehole revealed only very rare specimens which were

considered as of no biostratigraphic significance.

*Corresponding authors. Email: amr.daif@science.au.edu.eg; asdeaf75@yahoo.com

� 2013 AASP – The Palynological Society

Palynology, 2013

Vol. 00, No. 00, 1–27, http://dx.doi.org/10.1080/01916122.2013.828662

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Thus in most of the Egyptian palynological work,

palynostratigraphers have correlated different palyno-

morph assemblages with similar ones from other palaeo-

geographically related areas in order to date theEgyptian Cretaceous successions. As a result, several

informal, yet sometimes concept-incompatible (i.e., acme

versus total range) palynological zonal schemes (e.g., El-

Shamma 1991; Aboul Ela & Mahrous 1992) have been

proposed for different parts of the Cretaceous record of

Egypt. The informal zones proposed by Schrank and

Ibrahim (1995) represent the most complete palynologi-

cal zonal scheme for the Egyptian middle–upper Creta-ceous sedimentary sequence. Our zonal scheme uses

index taxa that have been recorded in the same phyto-

geographic province and have micropalaeontologically

calibrated age ranges. By using the same definitions as

Schrank and Ibrahim (1995), an attempt has been made

to develop a unified zonal scheme for the middle–upper

Cretaceous of the northernWestern Desert.

The goals of this project were to: (i) integrate lithos-tratigraphic and biostratigraphic schemes for the Abu

Tunis 1x borehole; (ii) propose an informal unified paly-

nozonal scheme for the middle–upper Cretaceous of the

northern Western Desert; (iii) correlate biostratigraphic

results with those from other palaeogeographically

related areas, such as the African and South American

phytogeographic provinces for sporomorphs, and the

Tethyan Realm for dinoflagellate cysts; (iv) provide thefirst quantitative description of the Egyptian Albian–

Cenomanian palynoflora, and (v) complete a taxonomic

study of all of the recorded taxa, especially of the strati-

graphically significant forms.

2. Geological setting and lithostratigraphic history

The north Western Desert is a vast area representing

the western part of a tectonically active structural

province referred to as ‘the unstable shelf’ by Said

(1962; Figure 1 Q1). Tectonism has played a major role in

the development and closure of several small rift basins

in this area. The oldest of these basins was developedin the late Palaeozoic, most probably during Late

Permian times, but most of these basins developed

around the earliest Cretaceous (Guiraud 1998;

Guiraud et al. 2001), with late Santonian folding caus-

ing inversion in most of the basins (Guiraud 1998).

The Western Desert is characterized by a northward

dipping plain comprising Eocene and Miocene

Figure 1. Simplified structural map of Egypt showing the location of the Abu Tunis 1x borehole (modified after Kerdany andCherif, 1990).

B=w in print; colour online

2 A.S. Deaf et al.

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carbonates, covered by Pliocene and Quaternary sedi-

ments along the Egyptian Mediterranean coast and the

Nile Delta areas (Hantar 1990).

Deposition of the upper lower Cretaceous sedimentsoccurred during a regressive phase, and fluvial processes

deposited the Albian Kharita Formation, which domi-

nates a large part of the Western Desert (Said 1990).Q2 In

contrast, the upper Cretaceous sediments indicate a

major transgressive phase. During the late Cenoma-

nian, regional subsidence related to Neotethyan rifting

took place across the northern African margin, and a

marine transgression covered the entire northernAfrican plate. As a result, transitional fluvial to shallow

marine deposits of the Bahariya Formation accumu-

lated in the Western Desert. During the Turonian,

marine conditions generally persisted across most of the

Western Desert, where a thick carbonate succession of

the the Abu Roash Formation was deposited in the

extreme northern Western Desert (Said 1990), where

the Abu Tunis 1x borehole is located. However, the lateTuronian Laramide tectonic event caused uplift and

basin inversion in some of the basins in the northern

Western Desert (Said 1990), and probably affected at

least the central part of the Matruh Basin in which the

Abu Tunis 1x borehole is located. However, by Conia-

cian times most of the northern Western Desert became

inundated by another marine transgression, during

which carbonates of the Khoman ‘B’ Formation weredeposited (Said 1990). By the late Santonian, right lat-

eral movement between Africa and Laurasia resulted in

northwest-directed compressive forces across the north-

eastern margin of the African plate (Meshref 1990;

Guiraud et al. 2001). These compressive forces in turn

resulted in a series of N-W folds (i.e., the Syrian Arc

System) associated with thrust faults across the north-

ern Western Desert (Meshref 1990; Guiraud et al.2001). As a result, the Santonian witnessed another

regression; continuing deposition of the lower Khoman

‘B’ Formation in the extreme north gave way south-

ward to marine clastic sedimentation across theWestern

Desert. During Campanian–Maastrichtian time, exten-

sion and subsidence dominated the northern Eastern

African margin (Guiraud & Bosworth 1999; Guiraud

et al. 2001), and most of the Western Desert was againcovered by deep marine waters, resulting in deposition

of a thick sequence of chalky limestone known as the

Khoman ‘A’ Formation (Said 1990).

3. Material and methods

Seventy-seven ditch cutting samples were collected

from between 7300 and 3500 ft (2225–1067 m) in theAbu Tunis 1x borehole. The borehole was drilled in the

Faghur area, at the centre of the Matruh Basin east of

the Faghur-Maamura High and north of the Umbarka

Subbasin: Lat. 31� 160 0800 N, and Long. 26� 500 4100 Eby WEPCO Q3(1968). A summary of the palynological

samples used with depths and total recovery of palyno-

morphs is provided in Appendix 1.Palynological processing involved acid digestion

using 36% hydrochloric acid (HCl) and 60% hydro-

fluoric acid (HF) (e.g., Phipps & Playford 1984; Green

2001). One tablet of a known quantity (12,542 grains/

tablet with V � 3.3%) of modern Lycopodium spores

(made by the Department of Quaternary Geology,

Lund University, batch no. 124961) was added, for

absolute abundance analysis, to each sample after thefirst HF decanting. Samples were boiled in 15 ml of

36% HCl for 1–2 minutes after being washed and

sieved through a 15-mm mesh to remove neo-formed

fluorides. Two permanent microscope slides were pre-

pared using Elvacite 2044 as a mounting medium.

Slides, residues and samples are stored in the Geologi-

cal Museum, Geology Department, Faculty of Science,

Assiut University, Egypt.Processed samples yielded well-preserved and

diverse assemblages of miospore-dominated palyno-

morphs and less frequent occurrences of lower-diver-

sity dinoflagellate cysts. Qualitative analysis was

carried out using an Olympus (BX41) transmitted-light

microscope (serial no. 8B25715) equipped with an

Infinity-1 digital camera for imaging. An alphabetical

list of the palynomorphs recorded in the Abu Tunis 1xis shown in Appendix 2. Quantitative analysis (grains/

gram) was determined by applying in the formula of

Stockmarr (1971) Q4as follows:

c ¼ Sc � Lt � t

Lc � wð1Þ

wherec ¼ concentration ¼ total number of specimens/

gram dried sediments

Sc ¼ number of specimens counted

Lt ¼ number of Lycopodium spores/tablet

t ¼ number of tablets added to the sample

Lc ¼ number of Lycopodium spores counted

w ¼ weight of dried sediments (g)

The first 250 grains were chosen for count as they pro-

vide a total maximum error of 7% in grain concentra-

tion according to the curve of Stockmarr (1971). To

detect any rare species, a further scan of the rest of

each mounted sample residue was also made.

4. Palynostratigraphy

4.1. Introductory comments

A careful review of Egyptian palynological literature

has shown that some of the previously proposed age

Palynology 3

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assignments have been misinterpreted because they

refer to species ranges, which have no independent age

control, or to sedimentary sequences of doubtful ages

or to processing cuttings with their inevitable cavingseffect. Therefore, dating of the Abu Tunis 1x borehole

was accomplished using independently calibrated

age ranges of index taxa compiled from the literature

(Figure 2), in addition to correlation with accurately-

dated contemporaneous regional and interregional

palynofloral assemblages. Correlation with interconti-

nental palynofloral assemblages was made in

the context of the Albian–Cenomanian ElateratesPhytogeographic Province of Herngreen et al. (1996).

In terms of the dinoflagellate cyst biostratigraphy,

it is important to note that many of the independently

calibrated events in the Albian of the European Tethys

(Davey & Verdier 1973; Habib & Drugg 1983; Erba

et al. 1999; Torricelli 2006) cannot be recognized in the

southern Tethyan region (e.g., Libya and Egypt), pos-

sibly due to environmental exclusion.Schrank and Ibrahim (1995) argued for early late

Cretaceous (Turonian–Coniacian) diachroneity of the

biostratigraphic units in terms of their concept and char-

acteristics in northeast Africa andWest Africa. This was

based on these authors extending the range of Droseri-

dites senonicus down into the late Turonian of Egypt

based on independent foraminifera-based dating. Whilst

the biostratigraphic units are defined differently (i.e.,acme zone versus total range), the palynological species

stratigraphic ranges are not at variance. This late Turo-

nian lower limit of Droseridites senonicus has also been

recorded in the foraminifera-dated late Turonian of

northeast Nigeria (Lawal & Moullade 1986). The inter-

regional correlation of biostratigraphic ranges and/or

successive events of selected index angiosperm (e.g.,

Cretacaeiporites) and gymnosperm (e.g., Droseridites

senonicus) pollen between the foraminifera-dated Ceno-

manian–Coniacian sedimentary sequence of North and

West Africa (Senegal and Ivory Coast: Jardin�e &

Magloire 1965; Egypt: Schrank & Ibrahim 1995;

Ibrahim 1996) shows a close match between these two

regions (Figure 3) and does not support the proposed

stratigraphic diachroneity. A greater similarity between

the palynofloral assemblage compositions and species’stratigraphic ranges could be proposed in the light of

African plate movement during the Cretaceous. This

tectonic plate was moving anticlockwise towards Laura-

sia, as a result of the opening of the southern Atlantic

Ocean, bringing formerly tropical regions into a more

subtropical position by the late Cretaceous (Figure 4).

Despite the fact that Morocco is confined to the

Albian–Cenomanian Elaterate Phytogeographic Prov-ince, however, some differences in the palynofloral

assemblages have been noted. In the Agadir-Essaouira

Basin and in the southwestern Tarfaya Basin, the

Albian–Cenomanian palynoflora exhibits Albian–

Cenomanian Elaterate Province characteristics, but

contains some temperate pollen grains, for example

Alisporites, Podocarpidites, Vitreisporites and Cerebro-

pollenites (Bettar & M�eon 2001, 2006) which are char-

acteristic of the Cerebropollenites Province of

Herngreen et al. (1996). Therefore, the micropalaeon-

tologically dated palynological work carried out in

West Africa (Jardin�e and Magloire, 1965: Senegal and

the Ivory Coast; Boltenhagen 1980: Gabon; Lawal &

Moullade 1986: Nigeria) which shows pure Albian–

Cenomanian Elaterate Province characteristics will beemployed here for assigning dates in the mid and late

Cretaceous sample interval. The local, partly foramin-

iferally controlled post-Cenomanian palynological

work of Schrank and Ibrahim (1995) will also be

followed.

The spore and pollen grain ranges have been used

for biostratigraphic purposes, with dinoflagellate cysts

providing additional supporting evidence. Dinoflagel-late cyst taxa have been found to be facies-controlled

in the middle Cretaceous sediments (Thusu et al.

1988), and additionally they are extremely diluted by

an abundance of microforaminiferal test linings in the

upper Cretaceous carbonate sediments. However, the

dinoflagellate cysts are the only biostratigraphic tools

that can be used to date the upper part of the carbonate

succession. Biozonation and age assessment of thestudied samples was based on the vertical quantitative

distributions by highest occurrence (HO) of all taxa

recovered (Figure 5). Lowest occurrences (LO) of

index sporomorphs were used to define biostrati-

graphic units where possible, but HO were used when

suitable LO were lacking. The numbers given in paren-

theses after the names of the taxa refer to the position

of these species in the quantitative range chart(Figure 5).

4.2. Palynological zonation (PZ) and age assessments

PZ1: Afropollis jardinus-Tricolporopollenites-

Elaterosporites klaszii Assemblage Zone (early–mid

Albian)

Samples: This zone includes samples 1 to 28, and spansa depth from 7300 to 5950 ft (2225–1814 m).

Definition: From the LO of Afropollis jardinus (36) to

just below the LO of Sofrepites legouxae (26) and Ela-

terosporites verrucatus (27).

Associated taxa: Verrucosisporites obscurilaesuratus (8),

Galeacornea causea (31), Ephedripites irregularis (33),

Rousea delicipollis (44), Triporites sp. (46), Retimonocol-

pites textus (49), Stellatopollis densiornatus (62),Afropol-lis aff. jardinus (66), Tetracolpites sp. (67), Senegalinium

aenigmaticum (85), Trichodinium castanea (87), Coroni-

fera oceanica (97), Oligosphaeridium albertense (105),

4 A.S. Deaf et al.

TPAL_A_828662.3d (TPAL) 21-08-2013 19:33

Figure 2. Compilation of the biostratigraphic range of most of the important Cretaceous marker species in different phytogeo-graphic provinces of North andWest Africa and north South America and the Tethyan Realm. Sources for African ranges: Doyleet al. 1982 (1); Hochuli 1981 (4, 11, 12); Hochuli and Kelts 1980 (1); Jan du Chene et al. (1978) (22, 25); Jardin�e 1967 (4, 5, 6, 8, 9,10, 11, 12, 14); Jardin�e and Magloire 1965 (1, 3, 4, 5, 6, 7, 8, 9, 13, 14, 15, 16, 17, 19, 21, 26); Lawal and Moullade 1986 (6, 7, 13,14, 15, 16, 17, 18, 22, 23, 24, 25, 26); Schrank and Ibrahim 1995 (2, 4, 6, 10, 11, 13, 16, 17, 20); (Tea-Yassi et al.) 1999 (18, 22, 23,24); Thusu and Van Der Eem (1985) (13). Sources for NS American ranges: Brenner 1968 (4, 6, 12); Herngreen 1973 (2, 3, 4, 5, 6,7, 8, 10, 11, 12, 13, 14); Herngreen 1975 (10); Herngreen and Due~naz Jimenez 1990 (12, 13, 14, 17, 19, 20, 21); Muller 1966 (1, 4,5, 12); Regali and Viana 1989 (2, 6); Regali et al. 1974 (1, 3, 5, 26). Sources for Tethyan ranges: Clark and Verdier 1967 (28, 30,31); Jan Du Ch�ene et al. 1978 (33); Roncaglia and Corradini 1997 (34); Schrank and Ibrahim 1995 (28, 29, 30, 31, 32, 33, 34);Torricelli and Amore 2003 (30, 31, 32, 34).

Palynology 5

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Oligosphaeridium complex (106), Oligosphaeridium

poculum (107), Pseudoceratium anaphrissum (111),Pseudoceratium securigerum (112).

Remarks: The microflora of this interval is character-

ized by the first appearance of Afropollis and elaterate

pollen grains (Figure 5; Plate 1). Tricolporate (e.g.,

Tricolporopollenites) and triporate (e.g., Triporites)

angiosperm pollen also first appeared in this interval

(Figure 5; Plate 2), whilst the tricolpate pollen (e.g.,

Tricolpites) present in the Aptian sediments of thisborehole are still present (Deaf 2009). Pteridophyte

spores and gymnosperm pollen grains dominate the

microfloral assemblage. Few of the angiosperm pollen

taxa which are present in the late Barremian are found

in the interval (Deaf 2009). The phytoplankton assem-

blage is more abundant than in the underlying interval,

rising from a maximum 1170 and average 105 cysts/

gram to a maximum 1750 and average 130 cysts/gram,but with decreasing species diversity (from 25 species

in the underlying interval to 12 species).

Discussion concerning age assessment: WEPCO (1968)

did not determine an age for the samples encompassed

by this palynozone, instead marking the succession

with a ‘no information’ label. The present study defines

an early Albian age for the base of this zone using the

LO of Afropollis jardinus (Figure 5), which occurs justabove the HO of the two marker Aptian forms Afro-

pollis operculatus and Afropollis zonatus (Figure 6).

The extinction of the latter two species was used by

both Doyle et al. (1982) and Schrank and Ibrahim

(1995) to document the Aptian/Albian boundary inWest Africa and Egypt respectively. Afropollis jardinus

is widely accepted as entering the stratigraphic record

in the early Albian in the Elaterates Phytogeographic

Province (Figure 2). In West Africa, it has been

recorded from foraminifera-dated rocks of early

Albian age in Senegal (as S. CI. 156 Incertae sedis) by

Jardin�e and Magloire (1965), and in Gabon-Congo-

Senegal by Doyle et al. (1982). In northern SouthAmerica, this same taxon has been recorded from fora-

minifera-dated sediments of early Albian age: in Brazil

(Herngreen 1973, 1975; Regali et al. 1974; Regali &

Viana 1989), Peru (Brenner 1968), and in Colombia

from sediments dated by ammonites as being of late

Albian–early Cenomanian age (Herngreen & Jimenez

1990). Finally, from western North Atlantic DSDP

Site 418A, Afropollis jardinus was also recorded (asReticulatasporites jardinus Type 3) from foraminifera-

dated rocks of late Albian–early Cenomanian age

(Hochuli & Kelts 1980).

The early–mid Albian boundary can be distin-

guished within this zone by the LO of Elaterosporites

klaszii, which is widely accepted to document the base

of the mid Albian in the Elaterates Phytogeographic

Province (Figure 2). This taxon has been recorded inSenegal and the Ivory Coast from foraminifera-dated

rocks of mid Albian–mid Cenomanian age (Jardin�e &

Magloire 1965; Jardin�e 1967). Foraminifera-dated

Figure 3. Cretaceous palynological biozonation correlation in North and West Africa and Northern South America (modifiedafter Schrank, 1992) and the palynological zonation proposed in the present study.

6 A.S. Deaf et al.

TPAL_A_828662.3d (TPAL) 21-08-2013 19:33

sediments from Brazil and Columbia have also indi-cated this taxon to be of mid Albian–mid Cenomanian

age (M€uller 1966; Herngreen 1973; Herngreen & Jime-

nez 1990), and it has also been recorded in the Albian–

Cenomanian of Peru (Brenner 1968). In northern Italy,

Elaterosporites klaszii has also been documented from

foraminifera-dated late Albian rocks by Hochuli (1981).

Although the index gymnosperm pollen Galeacor-

nea causea has been recorded from foraminifera-datedearly Cenomanian age in Senegal (e.g., Jardin�e 1967),

palynologically dated studies have extended its range

into the late Albian in Gabon and northeast Nigeria

(Doukaga 1980; Lawal & Moullade 1986). Galeacornea

causea was also recovered from rocks dated by forami-

nifera and ammonites as being of mid Albian–mid

Cenomanian age in Brazil and Colombia (M€uller 1966;Herngreen 1973; Herngreen & Jimenez 1990). Thus,

samples 27 and 28 in Abu Tunis 1x are likely to be of

mid Albian age.

Gnetaceaepollenites cf. clathratus occurs in the top-

most samples of this zone, and was identified in Sene-

gal by Stover (1963) from sediments of supposed

Cenomanian–Turonian age, and was later recorded

from the late Albian–mid Cenomanian of northeastNigeria (Lawal & Moullade 1986).

The occurrence of the gymnospermous tetrad

Droseridites senonicus (Plate 2) in samples 20 and

25 (19 and 23 grains/gram respectively) is problematic

because this species has only been recorded from rocks

of post-mid Albian age. For example, it is known from

385

390

395

400

405

410

Figure 4. World palaeogeographic maps showing the position of north Egypt during Albian and Turonian times (after Lawveret al. 2004).

B=w in print; colour online

Palynology 7

TPAL_A_828662.3d (TPAL) 21-08-2013 19:33

foraminifera-controlled Coniacian–Santonian sequen-

ces in northeast Nigeria and the Angola Basin

(Morgan 1978; Lawal & Moullade 1986), and in Egypt

from rocks also dated by foraminifera as of lateTuronian–early Santonian age (Schrank & Ibrahim

1995). The older occurrences of this taxon in Abu

Tunis 1x may be due to caving.

Based on the data above, an early to mid Albian

age is justified for this sample interval. Taken

together, the early–mid Albian age and dominantly

sandstone lithology of samples 1 to 28 suggests thatthese samples represent the lower Kharita Formation

(Figure 6), a clastic Albian–Cenomanian unit com-

posed of fine- to coarse-grained sandstones with

415

420

425

Figure 5. Quantitative (grains/gram) distribution chart by highest appearance of the recovered palynomorphs of the Abu Tunis1x borehole.

8 A.S. Deaf et al.

TPAL_A_828662.3d (TPAL) 21-08-2013 19:34

subordinate shale and carbonate interbeds in the

north Western Desert (Hantar 1990; Kerdany &

Cherif 1990).Correlation: This zone resembles Sequence X and XI

(early–mid Albian) identified by Jardin�e and Magloire

(1965) in the Senegal Basin, and corresponds exactly to

Zone I (early–mid Albian) of Herngreen (1973) in the

Maranh~ao Basin, Brazil. There are similar zones

defined in the northern Western Desert of Egypt: Zone

I of Sultan and Aly (1986), in the WD-9-15-1 well and

Zones II–III of Schrank and Ibrahim (1995), in theKahraman-1 well. Zone 2 (mid Albian) of Ibrahim

(1996), in the Ghazalat-1 well, corresponds to the

upper part of the current zone. In contrast, the Albian

Zone PSII of Mahmoud and Moawad (2002) in the

Sanhur-1X well very possibly corresponds only to thelower part of the current zone (i.e., early Albian),

because it lacks the mid Albian index form Elaterospor-

ites klaszii.

PZ2: Elaterosporites verrucatus-Sofrepites legouxae-

Cretacaeiporites Assemblage Zone (late Albian–early

Cenomanian)

Samples: This zone includes samples 29 to 39, which

cover a depth from 5900 to 5050 ft (1798–1539 m).

430

435

440

445

Figure 5. Continued

B=w in print; colour online

Palynology 9

TPAL_A_828662.3d (TPAL) 21-08-2013 19:34

Definition: From the LO of Sofrepites legouxae (26)

and Elaterosporites verrucatus (27) to HO of Sofrepiteslegouxae (just above the HO of Elaterosporites

verrucatus).

Associated taxa: Classopollis classoides (19), Ephedri-

pites spp. (20), Araucariacites australis (24), Afropollis

jardinus (36), Tetraporopollenites sp. (39), Rousea deli-

cipollis (44), Tricolpites vulgaris (48), Retimonocolpites

textus (49), Rousea brenneri (50), Tricolpites parvus

(51), Triporopollenites spp. (53), Tricolpites micromunus

(54), Dichastopollenites ghazalatensis (55), Tricolpites

cf. crassimurus (56), Tricolpites sagax (58),

Retimonocolpites ghazalii (59), Stephanocolpites sp.

(60), Stellatopollis barghoornii (61), Stellatopollis den-

siornatus (62), Striatopollis cf. trochuensis (63) Tricho-

dinium castanea (87), Xiphophoridium alatum (95),

Cribroperidinium edwardsii (108).

Remarks: The microfloral assemblage of this interval is

characterised by elaterate pollen grains and a notice-

able increase in abundance of the tricolpate angio-

sperm pollen. The genus Afropollis also shows a

significant increase from a maximum 1500 (average�400) grains/gram in the underlying interval to a maxi-

mum of 5375 (average �3400) grains/gram. This

450

455

460

465

470

Figure 5. Continued

B=w in print; colour online

10 A.S. Deaf et al.

TPAL_A_828662.3d (TPAL) 21-08-2013 19:34

Plate 1. The sample/slide number, England Finder coordinates, and position of taxa on the quantitative range chart (numbers inparentheses) are indicated for all specimens; scale bar represents 20 mm. The microphotographs were all taken using plain trans-mitted light.

B=w in print; colour online

Palynology 11

TPAL_A_828662.3d (TPAL) 21-08-2013 19:35

Plate 2.

B=w in print; colour online

12 A.S. Deaf et al.

TPAL_A_828662.3d (TPAL) 21-08-2013 19:35

increase is accompanied by a decrease in diversity of

spore species, which are mainly represented by Deltoi-

dospora and Cicatricosisporites. The abundance of phy-

toplankton continues to decrease (maximum 244,average 100 cysts/gram) throughout the interval, but

with a similar low diversity (�14 species) to the under-

lying interval.

Discussion concerning age assessment: Sofrepites

legouxae is an index taxon in this interval (Figure 5;

Plate 1), which was found to range from the late Albian

to early Cenomanian in foraminifera-dated rocks of

Senegal by Jardin�e and Magloire (1965), Jardin�e(1967), and in Brazil by Herngreen (1973) and

Herngreen and Jimenez (1990). Elaterosporites

verrucatus is recorded in Senegal and the Ivory Coast

from foraminifera-dated mid Albian–early Cenoma-

nian sediments (Jardin�e & Magloire 1965; Jardin�e1967: Figure 2). However, in foraminifera- and ammo-nite-dated rocks from Brazil, this same taxon was

recorded from the latest mid to earliest late Albian

(Herngreen 1973; Regali et al. 1974; Herngreen &

Jimenez 1990), and a single occurrence has been

reported from the middle part of the ammonite-dated

late Albian of Columbia (Herngreen & Jimenez 1990).

Cretacaeiporites densimurus appears in the lower part

of this sample interval, and was first described bySchrank and Ibrahim (1995) from foraminifera-dated

rocks of early–mid Cenomanian age, and was later

475

480

485

490

495

500

Figure 6. The Abu Tunis 1x borehole with lithological column, sample positions, original age dating, key biostratigraphic eventsand ages deduced in the current work.

B=w in print; colour online

Palynology 13

TPAL_A_828662.3d (TPAL) 21-08-2013 19:35

recorded by Ibrahim (2002) from the late Albian–early

Cenomanian of Egypt. Single specimens of other spe-

cies of this genus have been found in the upper part of

this interval: Cretacaeiporites polygonalis, which has alate Albian–late Cenomanian range in Senegal (Jardin�e& Magloire 1965), and Cretacaeiporites mulleri, which

ranges from the late Albian up to the Santonian in Sen-

egal and northeast Nigeria (Jardin�e & Magloire 1965;

Lawal & Moullade 1986). In Brazil, Cretacaeiporites

mulleri was recorded as of late Albian–mid Cenoma-

nian age (Herngreen 1973).

Appearing throughout the upper part of this inter-val, the range of Elaterocolpites castelainii was used to

document the base of the late Albian and top of the mid

Cenomanian in the Elaterates Province in Senegal

(Jardin�e & Magloire 1965; Jardin�e 1967: FigureQ5 ) and

Brazil (Herngreen 1973; Herngreen & Jimenez 1990).

This taxon was also recorded by Hochuli (1981)

from the foraminifera-dated late Albian of southern

Switzerland. Other late Albian–early Cenomanianelaterate taxa present in this zone are Elaterosporites

acuminatus and Elaterosporites protensus (Figure 6).

Elaterosporites acuminatus was reported from the late

Albian–early Cenomanian of Senegal (Jardin�e 1967),

and occurs in the lower part of this interval. Elaterospor-

ites protensus is found in the same samples and was

recorded from foraminifera-dated rocks of mid Albian

to latest Albian/earliest Cenomanian age in Senegal, theIvory Coast (Jardin�e & Magloire 1965; Jardin�e 1967)

and Brazil (Herngreen 1973; Herngreen & Jimenez,

1990).

Afropollis kahramanensis was recorded by Schrank

and Ibrahim (1995) and Ibrahim (2002) from the fora-

minifera-dated early–mid Cenomanian of Egypt. The

former authors mentioned that the taxon described as

Pollen PO-304 by Lawal and Moullade (1986) from thesupposed late Albian–mid Cenomanian of northeast

Nigeria was identical to their new species; therefore, the

presence of Afropollis kahramanensis in this

interval does not contradict the proposed late Albian–

Cenomanian age. Foveotricolpites gigantoreticulatus

was recorded from foraminifera-dated rocks of

Turonian–Santonian age in Senegal and northeast

Nigeria (Jardin�e & Magloire 1965; Lawal & Moullade1986). However, Schrank and Ibrahim (1995) and

Schrank and Mahmoud (1998) documented an older

occurrence of this species in the palynologically dated

Albian–Cenomanian of Egypt. Thus, the presence of

Foveotricolpites gigantoreticulatus in the middle part of

this interval may not conflict with the proposed

late Albian–early Cenomanian age of the interval.

The rare presence of Triporites spp. in this intervalis in accordance with the late Albian–early Cenomanian,

as rare Triporites spp. were recorded from the early

Cenomanian of Senegal (Jardin�e &Magloire 1965).

The questionable occurrence of Droseridites bacu-

lites in this interval, which was described by Ibrahim

(1996) from the palynologically dated lower Turonian

of Egypt, may be attributed to caving.As for the dinocysts, Florentinia berran appears in

the lower part of the interval, a taxon that was recorded

from the Albian–early Cenomanian of the southern

Tethyan Realm in Morocco and northeast Libya

(Below 1982, 1984; Uwins & Batten 1988). Florentinia

laciniata and Florentinia mantellii first appear in this

interval and are known from ammonite-dated Aptian–

early Cenomanian sequences in the Tethyan Realm:they continue upward into the overlying intervals.

Based on the presence of the rare late Albian–early

Cenomanian elaterates and the marker angiosperm

forms mentioned above, a late Albian–early Cenoma-

nian age is postulated for the interval.

Samples from this part of Abu Tunis 1x were recog-

nized by WEPCO (1968) as ‘Cenomanian Clastics’.

However, the dominant sandstone composition of sam-ples 29–39 when combined with the late Albian–early

Cenomanian age proposed here suggests allocation of

these sediments to the upper Kharita Formation.

Correlation: The palynomorph assemblage recorded in

this zone is identical to that described in Zone II (late

Albian–early Cenomanian) of Herngreen (1973) from

the Maranh~ao Basin, Brazil. Similar late Albian–early

Cenomanian assemblages that share some of the elater-ate and angiosperm pollen grains (but not Sofrepites

legouxae) are those of sequences IX of Jardin�e and

Magloire (1965) in the Senegal Basin, and Subzone Ia

of Lawal and Moullade (1986) in the upper Benue

Basin, northeast Nigeria. The same applies to the

assemblage of intervals 3–4 in the Manndra 1 well and

intervals c–d in the Algerian Oued Melah 1 well of

Foucher et al. (1994). Similar late Albian–earlyCenomanian zones (but also lacking Sofrepites

legouxae) have also been documented from the Egyp-

tian northern Western Desert, for example, Zone III of

Aboul Ela and Mahrous (1992) in the East Tiba-1 well,

Zone IV of Schrank and Ibrahim (1995), in the

Kahraman-1 well, and assemblage ‘A’ of Ibrahim

(2002), in the Abu Gharadig-5 well. The upper part of

Zone II (mid/late Albian–early Cenomanian) of El-Beialy et al. (2011) in the Gebel Rissu-1 well is very

similar to the current palynozone, and this time

S. legouxae was recorded.

PZ3: Sofrepites legouxae Partial Range Zone (early–

?mid Cenomanian)

Samples: This zone spans samples 40 to 46 and a depthfrom 4950 to 4650 ft (1509–1417 m).

Definition: From the HO of Sofrepites legouxae (26) to

the LO of Proteacidites cf. africaensis (35).

505

510

515

520

525

530

535

540

545

550

555

560

565

570

575

580

585

590

595

600

605

14 A.S. Deaf et al.

TPAL_A_828662.3d (TPAL) 21-08-2013 19:35

Associated taxa: Deltoidospora spp. (1), Crybelosporites

pannuceus (2), Cicatricosisporites orbiculatus (4), Bal-

meiopsis limbatus (18), Classopollis classoides (19),

Ephedripites spp. (20), Elaterosporites klaszii (21),Araucariacites australis (24), Elaterocolpites castelainii

(25), Afropollis jardinus (36), Rousea delicipollis (44),

Cretacaeiporites mullerii (45), Triporites spp. (46), Rou-

sea cf. miculipollis (47), Florentinia mantellii (99), Flor-

entinia laciniata (103).

Remarks: Sporomorphs are represented by pterido-

phyte spores, pollen from xerophytic gymnosperms,

two elaterate, and five angiosperm pollen species. Thephytoplankton assemblage shows a slight increase in

abundance over the interval below (maximum 370,

average 120 cysts/gram) but with a consistently low

diversity (�10 species), and provides an important

characteristic for dating the interval.

Discussion concerning age assessment: Proteacidites cf.

africaensis, which is widely accepted to document the

base of the mid Cenomanian in the western part ofthe Elaterates Province (e.g., Jardin�e & Magloire

1965; Lawal & Moullade 1986) first appears in

the overlying interval (in Sample 47), and thus

delineates the lower boundary of the mid

Cenomanian. Consequently, the present interval may

be of early Cenomanian age.

Florentinia berran, which was found in sediments as

young as the early–mid Cenomanian in Egypt(Schrank & Ibrahim 1995; Ibrahim 2002), becomes

extinct in the lower part of the overlying interval, thus

favoring an early–mid Cenomanian age for this sample

interval. Interestingly, an exact match of the biostrati-

graphic events recorded herein: the highest occurrence

of Elaterocolpites castelainii, the occurrence of Elatero-

sporites klaszii, Afropollis jardinus, and rare triporate

pollen grains, with those recorded in the Zone V(early–?mid Cenomanian) of Schrank and Ibrahim

(1995) also supports an early–?mid Cenomanian age

for the current zone.

WEPCO (1968) referred to the interval from which

samples 40 to 46 come as ‘Cenomanian Clastics’. The

mixed, clastic-carbonate lithology of these samples

and the presumed early–?mid Cenomanian age indi-

cates this part of the sequence can now be identifiedas the lower Bahariya Formation (Figure 6), as the

lower part of this formation in the north Western

Desert is composed of sandstones with alternating

with shales and frequent limestone horizons. This

formation conformably overlies the Kharita

Formation and underlies the Abu Roash Formation

(Norton 1967).

Correlation: Regional comparisons of Egyptian (early–mid Cenomanian) palynozones reveal that Zone V of

Schrank and Ibrahim (1995) in the Kahraman-1 and

Abu Gharadiq-18 wells, Zone 3 of Ibrahim (1996) in

the Ghazalat-1 well and assemblage ‘B’ of Ibrahim

(2002) in the Abu Gharadiq-5 well can be correlated

with the present zone.

PZ4: Proteacidites cf. africaensis Total Range Zone(mid–late Cenomanian)

Samples: This zone includes samples 47 to 62 and cov-

ers a depth from 4600 to 3850 ft (1402–1173 m).

Definition: Total range of Proteacidites cf. africaensis

(35).

Associated taxa: Deltoidospora spp. (1), Crybelosporitespannuceus (2), Alisporites cf. grandis (17), Balmeiopsis

limbatus (18), Ephedripites spp. (20), Elaterosporites

klaszii (21), Retimonocolpites variplicatus (41), Tricol-

poropollenites sp. (43), Senegalinium aenigmaticum

(85), Trichodinium castanea (87), Surculosphaeridium

cf. longifurcatum (88), Xiphophoridium alatum (95),

Florentinia spp. (94), Florentinia mantellii (99).

Remarks: The palynofloral assemblage of this intervalshows characteristics similar to the underlying interval,

but with a continuing decrease in the abundance and

diversity of spores (maximum 48/average 24 grains/

gram, and only two species), the complete disappear-

ance of gymnosperm pollen grains at the end of the

interval, and the occurrence of nine angiosperm pollen

taxa. Phytoplankton increase in abundance (maximum

3160, average 140 cysts/gram), but have a very low spe-cies diversity (�8 species).

Discussion concerning age assessment: The inception of

Proteacidites cf. africaensis at the base of this interval

is taken here to mark the lower boundary of the mid

Cenomanian interval, as it was recorded in West

Africa, in Senegal and Nigeria (Jardin�e & Magloire

1965; Jardin�e 1967; Lawal & Moullade 1986) and

Brazil (Herngreen 1973), from foraminifera-datedrocks of mid–late Cenomanian age.

The occurrence of Afropollis jardinus in these sam-

ples requires explanation. The extinction of this species

has been shown to be diachronous across palaeotropi-

cal African regions, occurring either in the early or mid

Cenomanian, a character attributed by Doyle et al.

(1982) to palaeoclimatic influences. In the foraminif-

era-dated Gabon reference section for Afropollis spe-cies, Doyle et al. (1982) found that the abundance of

Afropollis jardinus declined in the early Cenomanian

Subzones C-XIIb and C-XIIc, and disappeared before

the appearance of Proteacidites africaensis (as Triorites

africaensis) in mid-late Cenomanian age sediments

(Jardin�e & Magloire 1965; Jardin�e 1967). However, in

Senegal Doyle et al. (1982) noted that Afropollis jardi-

nus diminishes in the early mid Cenomanian – laterthan in the Gabon section – and disappeared by the

end of the mid Cenomanian, contemporary with the

appearance of Proteacidites africaensis. Doyle et al.

610

615

620

625

630

635

640

645

650

655

660

665

670

675

680

685

690

695

700

705

710

715

Palynology 15

TPAL_A_828662.3d (TPAL) 21-08-2013 19:35

(1982) interpreted this persistence of Afropollis

jardinus through the mid Cenomanian as due to more

favourable (wetter) conditions. An interpretation that

was later supported by Schrank (2001)Q6 as he suggestedAfropollis thrived in humid coastal areas. The same

scenario could also apply to Egypt, which was at a

palaeolatitude similar to those of Senegal and

where Afropollis jardinus may have persisted into the

mid Cenomanian due to local humid coastal

conditions.

The HO of Afropollis jardinus is found at the top of

both the foraminifera-dated Zone V of Schrank andIbrahim (1995) and Assemblage Zone ‘B’ of Ibrahim

(2002), both of which are known to be of early–mid

Cenomanian age. Therefore, the HO of Afropollis jar-

dinus in sample 53 in Abu Tunis 1x is used to delineate

the upper boundary of the mid Cenomanian. Afropollis

kahramanensis, which was recorded in the foraminif-

era-dated lower–middle Cenomanian rocks of Egypt

(Schrank & Ibrahim 1995; Ibrahim 2002), has its HOat the same level as the HO of Afropollis jardinus. Ela-

terosporites klaszii has its highest occurrence in the

lower part of this interval, and is known to terminate

in the mid Cenomanian in the Elaterates Province

(Jardin�e & Magloire 1965; M€uller 1966; Jardin�e 1967;

Herngreen 1973; Herngreen & Jimenez 1990: Figure 2).

The presence of Cretacaeiporites densimurus is also

consistent, as it has its uppermost occurrence at thetop of the mid Cenomanian in Egypt (Schrank &

Ibrahim 1995; Ibrahim 2002). The late Cenomanian

age of the upper part of this zone is inferred from the

upward continuation of Proteacidites cf. africaensis

and from the very rare occurrence (and later complete

disappearance) of the gymnosperm pollen Classopollis

spp.. In Senegal, Jardin�e and Magloire (1965) recorded

high abundances of Classopollis spp. (up to 80%) fromthe Barremian to the mid Cenomanian, which then

declined rapidly and became extinct by the end of the

late Cenomanian. Similarly, Schrank and Ibrahim

(1995) and Ibrahim (2002) recorded rare final occur-

rences of Classopollis in the middle Cenomanian rocks

of Egypt.

A mid–late Cenomanian age is therefore assigned

to this interval based on the presence of the indexgymnosperm and angiosperm taxa mentioned above.

The mid to late Cenomanian age combined with the

dominantly carbonate lithology of the samples in this

palynozone suggests the unit should be identified as

the upper Bahariya to lower Abu Roash formations.

WEPCO (1968) did not recognise such subdivisions

in their original study of Abu Tunis 1x, but the

new assignments are consistent with the known litho-logical and age characteristics of the Abu Roash

Formation, a limestone sequence with interbeds of

shale and sandstone known to range in age from late

Cenomanian to Turonian (Hantar 1990). The Abu

Roash is conformably underlain by the Bahariya

Formation and overlain by the Khoman ‘B’

Formation.Correlation: The characteristics of this palynozone

strongly resemble the mid–late Cenomanian Sequence

VII of Jardin�e and Magloire (1965) in the Senegal

Basin and the ‘Triorites’ africaensis Zone (II) of Lawal

and Moullade (1986), upper Benue Basin, northeast

Nigeria. Zone III (late Cenomanian) of Herngreen

(1973), Brazil, only equates to the upper part of the

present zone. Other zones similar to the current onehave been identified in the northern Western Desert:

Zone VI of Schrank and Ibrahim (1995) in the Kahra-

man-1 and Abu Gharadiq-18 wells, and Zone 4 of

Ibrahim (1996) in the Ghazalat-1 well, both zones hav-

ing been allocated a mid?–late Cenomanian age. Zone

I (late Cenomanian) of El Beialy et al. (2011) in the

Gebel Rissu-1 well could also correspond to the cur-

rent zone, as the latter authors have equated their zoneto the aforementioned Zone VI of Schrank and

Ibrahim (1995). This could be further supported by the

mid Cenomanian Elaterosporites klaszii and Cretacaei-

porites densimurus which occur in the current zone as

they do in the lower part (samples 3–5) of Zone I of El

Beialy et al. (2011).

PZ5: Canningia senonica Total Range Zone (early?

Santonian)

Samples: This zone includes samples 73 to 74 and cov-

ers a depth from 3250 to 3200 ft (991–975 m). The two

overlying samples, 75 and 76, are barren. Sample 77 is

separated from 75 and 76 by an unconformity surface

(identified in the well log by WEPCO 1968) and exhib-

its very poor dinoflagellate recovery and was deemedbarren (Figure 5).

Definition: Total range of Canningia senonica (79).

Associated taxa: Alisporites cf. grandis (17), Spiniferites

spp. (77), Exochosphaeridium bifidum (80), Dinogym-

nium spp. (82), Senegalinium aenigmaticum (85), Tri-

chodinium castanea (87), Surculosphaeridium cf.

longifurcatum (88), Downiesphaeridium sp. (92).

Remarks: Palynological assemblages from this zone aredominated by phytoplankton, with only one terrestrial

palynomorph being present: the gymnosperm Alispor-

ites cf. grandis.

Discussion concerning age assessment: Canningia senon-

ica (Plate 3) appears in both samples 73 and 74

(Figure 5), a species characteristic of the Santonian–

Maastrichtian. Canningia senonica has been recorded

from the ammonite-dated late Santonian of the Isle ofWight, southern England (Clarke & Verdier 1967),

from the foraminifera-dated early Santonian–late

Maastrichtian of Egypt (Schrank & Ibrahim 1995),

720

725

730

735

740

745

750

755

760

765

770

775

780

785

790

795

800

805

810

815

820

825

16 A.S. Deaf et al.

TPAL_A_828662.3d (TPAL) 21-08-2013 19:35

Plate 3.

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Palynology 17

TPAL_A_828662.3d (TPAL) 21-08-2013 19:35

and from the nannoplankton-dated late Campanian of

Italy (Torricelli & Amore 2003).

This interval contains a few long-ranging species

of pre-Coniacian to Maastrichtian age, such asChlamydophorella discreta which appears first in this

interval, a species which has its highest occurrence in

the late Santonian of the Isle of Wight (Clarke &

Verdier 1967), and ranges in Egypt from the early

Cenomanian to the Turonian (Schrank & Ibrahim

1995). Isabelidinium acuminatum was recorded (as

Deflandrea acuminata) from the ammonite-dated late

Santonian of the Isle of Wight (Clarke & Verdier1967), and from the foraminifera-dated Coniacian–

Santonian of Algeria (Foucher et al. 1994). Dinogym-

nium denticulatum, recorded from the ammonite-dated

Late Santonian of the Isle of Wight (Clarke & Verdier

1967), from the foraminifera-dated sequences of

Coniacian–Maastrichtian age in Egypt (Schrank &

Ibrahim 1995), and of Coniacian–Santonian age in

Algeria (Foucher et al. 1994). Eucladinium gamban-

gense, which was identified by Cookson and Eisenack

(1970) from the Senonian rocks of Australia, occurs

at the base of this interval.

There are no index angiosperm pollen taxa that one

might expect for the Turonian, such as Foveotricolpites

giganteus or Foveotricolpites gigantoreticulatus, which

are characteristic of the early Turonian–Santonian in

West Africa (e.g., Jardin�e & Magloire 1965; Lawal &Moullade 1986; Schrank & Ibrahim 1995). Nor is Dro-

seridites senonicus present, diagnostic of the late

Turonian–early Santonian (Lawal & Moullade 1986;

Salard-Cheboldaeff 1990; Schrank & Ibrahim 1995;

Ibrahim 1996).

Thus, based on the presence of the early Santonian

index form Canningia senonica and the complete

absence of index Turonian sporomorphs mentionedabove, an early Santonian age is proposed for this

interval. The WEPCO report (1968) did not allocate an

age to this part of the borehole succession.

The proposed ages and carbonate lithology of sam-

ples 73 and 74 suggest this part of the sequence can be

assigned to the lower Khoman ‘B’ Formation, which is

easily recognized in north Egypt by its white chalk/

chalky limestone lithology. This unit unconformablyoverlies the Abu Roash Formation in the Abu Tunis

1x borehole (as is the case in many basins of the north

Western Desert), and conformably underlies the

Khoman ‘A’ Formation. The Khoman ‘B’ Formation

has been allocated a Coniacian–Santonian age (Hantar

1990; Kerdany & Cherif 1990).

Correlation: The dinoflagellate Zone 8 (early

Santonian) of Schrank and Ibrahim (1995), in theKahraman-1 well in the northern Western Desert

of Egypt, is the only zone that correlates to the

palynozone 5.

5. Discussion of the Abu Tunis 1x palynostratigraphy

Semi-quantitative frequencies are closed scales in

which measured variables depend on each other, and

thus reflect a false picture of real quantitative distribu-tions of variables measured. Thus, this section deals

with the quantitative (grains/g of sediments) distribu-

tions but combined with semi-quantitative to explore

effect of data closure on semi-quantitative distribu-

tions, and to enable a wider comparison of quantitative

data with its counterparts (i.e., semi-quantitative) in

regional and interregional contexts.

5.1. Albian–Cenomanian Elaterates PhytogeographicProvince

5.1.1. Quantitative characteristics of the Albian–

Cenomanian palynoflora from the present study

The palynoflora of the Abu Tunis 1x exhibits a remark-

able similarity to that of the Albian–Cenomanian Ela-

terates Phytogeographic Province of Herngreen et al.

(1996). Features diagnostic for the province are repre-

sented here by the appearance and later increased

abundances of Afropollis jardinus (0.4–54%, avg. 15%;4–7003, avg. 1219 grains/g), and occurrence of Crybelo-

sporites (0.4–3.6%, avg. 2%; 4–523, avg. 118 grains/g).

Other typical criteria for the recognition of this prov-

ince found in the Abu Tunis 1x borehole are the

appearance and diversification of gymnospermous ela-

terate pollen grains: Elaterosporites (0.4–7.6%, avg.

39%; 4–854, avg. 262 grains/g), Elaterocolpites (0.4%;

40–52 and avg. 46 grains/g), Galeacornea (0.8%;26 grains/g), and Sofrepites(0.4–2.4%, avg. 1%; 21–465,

avg. 146 grains/g), and the presence of Cretacaeiporites

(0.4–1.6%, avg. 0.9%; 19–174, avg. 75 grains/g). The

diversification of Tricolpites (0.4–3.2%, avg. 1.6%; 25–

319, avg. 90 grains/g), Tricolporites (0.4%; 12–58, avg.

35 grains/g), and Triporites (0.4–1.2%, avg. 0.6%; 12–

119, avg. 43 grains/g) accompanied with a drop in

abundance of smooth trilete spores from (17–87%, avg.53%; 1332–40762, avg. 7663 grains/g) in the late Albian

to (0.8–38%, avg. 11; 9–2195, avg. 639 grains/g) in the

Cenomanian are important features. The complete

absence of bi- and tri-saccate gymnospermous pollen

grains, and the abundance (0.4–5.6%, avg. 0.2%; 5–

1394, avg. 270 grains/g) and later disappearance of

Classopollis in the mid–late Cenomanian are also diag-

nostic features.

5.1.2. Regional mid Albian–early Cenomanian acme of

Afropollis jardinus

The genus Afropollis jardinus, as mentioned in Section4.2.4, shows different biostratigraphic and semi-quan-

titative distributions between northeast Africa and

West Africa that have been interpreted to be the result

830

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18 A.S. Deaf et al.

TPAL_A_828662.3d (TPAL) 21-08-2013 19:35

of palaeoecological factors. Palynological studies of

this taxon in Egypt have so far been semi-quantitative,

and no attempt has been made to explore the possible

quantitative significance of the species on a regionalscale. Quantitatively, the temporal abundance of Afro-

pollis jardinus (Figure 7) shows very low concentra-

tions (� 33–738, avg. 350 grains/g; 0.4–17%, avg. 4%)

in the lower Albian of the Abu Tunis 1x well until the

middle Albian–lower Cenomanian marine sediments,

where a peak abundance is recorded of 1400–7003 and

average 2900 grains/g (24–62%, avg. 34%). In an adja-

cent basin, about 51 km to the southwest of AbuTunis, Schrank and Ibrahim (1995) recorded a simi-

larly high relative abundance acme (31–61%, avg. 40%)

in the mid Albian–early Cenomanian of the

Kahraman-1 well. The slightly higher percentages of

the species in the mid Albian of the Kahraman-1 well

may potentially be attributed to more favourablelithologies, as there is a greater shale content in this

part of the well when compared to Abu Tunis 1x. Simi-

larly, about 190 km to the southeast of Abu Tunis 1x,

the same mid Albian–early Cenomanian acme (11–

64%, avg. 34%) of Afropollis jardinus was recorded

from the Bed 1–1 well (El-Beialy 1994). Consistency

between quantitative and semi-quantitative values

through the Afropollis jardinus acme as recorded frommarine sediments of three wells in three different basins

supports the contention made by Schrank (2001) that

this acme event may have been controlled by locally

humid conditions associated with proximity to shallow

marine depositional environments, as the Afropollis-

parent plant is thought to have thrived in humid

coastal plans. Interregionally, similar mid Albian–early

Cenomanian (10–40%, avg. 14%) peaks of Afropollis

jardinus have been documented in Senegal (Doyle et al.

1982). In Gabon, the appearance of the Afropollis jar-

dinus acme in the early–mid Albian differs from those

recorded in Egypt and Senegal, and may be due to dif-

ferences in palaeogeographic position and geological

history of the basins (Doyle et al. 1982). In this light,

acmes of Afropollis are only of biostratigraphic rele-

vance on a regional – i.e., not interregional – scale.We believe that the acme event of Afropollis jardi-

nus in the north Western Desert of Egypt is an impor-

tant regional mid Albian–early Cenomanian bioevent.

5.2. Senonian palynoflora from the present study

The early Santonian palynoflora of the Abu Tunis 1x

borehole is exclusively represented by marine phyto-planktons and completely lacks any terrestrial palyno-

morphs that would be representative for the Senonian

Palmae Province of Herngreen et al. (1996).

5.3. Late Cretaceous uplift of the Faghur area

An unconformity surface is identified by WEPCO

(1968) in the composite log of the Abu Tunis 1x bore-hole separating the palynomorph-barren interval

assigned by WEPCO (1968) as of Turonian age from

the overlying dinoflagellate-identified lower Khoman

‘B’ (early? Santonian) Formation (Figure 6). This may

indicate a short time cessation in deposition and ero-

sion, which may be related to the Turonian uplift that

affected most of the Western Desert basins (e.g.,

Kerdany & Cherif 1990). By the early Santonian, theFaghur area had become marine, based on the marine

palynomorphs recorded in the Abu Tunis 1x borehole

sediments.

935

940

945

950

955

960

965

970

975

980

985

990

995

Figure 7. Quantitative and semi-quantitative vertical distri-butions of Afropollis jardinus in the Abu Tunis 1x boreholecompared to semi-quantitative distributions in Egyptianwells and Senegal and Gabon Albian–Cenomanian sections.

Palynology 19

TPAL_A_828662.3d (TPAL) 21-08-2013 19:36

5.4. A Unified mid-Late Cretaceous palynologicalzonal scheme for the northern Western Desert

The zonal scheme devised for the Abu Tunis 1x bore-

hole suggests the use of unified biozones of those of

Schrank and Ibrahim (1995) and our zones, which are

at least applicable to the Egyptian Albian–middle Cen-

omanian rocks of the northern Western Desert. This is

because the Albian–Cenomanian diagnostic taxa have

been widely recorded in Egypt and have calibrated

stratigraphic ranges across the Albian–CenomanianElaterates Phytogeographic Province (PZ1-4). There

are some difficulties when dealing with the upper Ceno-

manian rocks. This may be due to change in lithology

and depositional environment from clastic shallow

marine (as in the Kahraman-1 well, Schrank & Ibrahim

1995) to carbonate inner neritic environment towards

the north (as in the Abu Tunis 1x well). More difficul-

ties arise when dealing with Turonian and youngerrocks. Palynomorph recovery from post Cenomanian–

Maastrichtian carbonates is rather low due to deeper

neritic marine environment, regional late Cretaceous

tectonism and a global late Cretaceous decline in gen-

era and species diversity of palynomorphs. In order to

draw a complete and unified mid-late Cretaceous zonal

scheme for the Egyptian northern Western Desert, it is

suggested that further palynological (preferably agecontrolled) work on successions of adjacent and less

tectonically affected basins that were slightly close to

palaeovegetation cover is needed.

6. Systematic palynology

A. Elaterate pollen

Genus: Elaterosporites Jardin�e 1967Type species: Elaterosporites verrucatus (Jardin�e andMagloire) Jardin�e 1967, p. 244, pl. 2, figs. E-G, pl. 3,fig. G.Elaterosporites verrucatus (Jardin�e and Magloire)Jardin�e 1967Plate 1, figs. 6, 8

Description: Grains with ellipsoidal plano-convex to

sub-hemispherical body with three U-shaped horns

implanted on the convex distal face. Proximal face flator depressed, bounded by 3–8 mm wide annular ring

structure parallel to the equator ending with protrud-

ing ends parallel to the long axis. These three U-shaped

appendages with their middle part running parallel to

the longest axis; one is placed over the distal pole, the

other two are placed laterally. The appendages are

solid and of uniform diameter. Exine is granulate to

verrucate. Main body length 38–64 mm and breadth25–45 mm.

Dimension: Maximum main body length (2 specimens)

40 (52.5) 65 mm, breadth undetermined due to grain

orientation, annular ring 3–8 mm wide; appendage

length 33 (36.5) 40 mm, width 3.5 (4.5) 5.5 mm.

Remark: Elaterosporites verrucatus can be distin-

guished from Elaterosporites acuminatus (Plate 1, fig-ures 7, 14) by its 3–8 (here also 3–8) mm wide annular

ring structure and granulate to verrucate exine (here

verrucae 1–3.5 mm wide) versus 8–12 (here 6–8) mm

wide annular ring and densely and uniformly packed

exine with spines 4–6 (here 4–5) mm high and 2.5–3

(here 2.5–3.5) mm wide.

Elaterosporites protensus (Stover) Jardin�e 1967Plate 1, fig. 9

Remark: Stover (1963) distinguished Elaterosporites

protensus from Elaterosporites acuminatus by its larger

size (Elaterosporites acuminatus: 52 � 28 mm), greater

ring width, and appendages ending with round tips

rather than with sharply tipped ends as in the latterspecies. Here dimensions of Elaterosporites protensus

are: maximum main body length (1 specimen) 65 mm,

breadth 35 mm, annular ring 7–9.5 mm wide; append-

age length and width undetermined due to grain orien-

tation. Dimensions of Elaterosporites acuminatus are:

maximum main body length (2 specimens) 39 (44)

49 mm, breadth 27 (28) 29 mm, annular ring 6–8 mm

wide; appendage length 32 (36) 40 mm.

B. Pollen tetrad

Genus: Droseridites Cookson 1947 ex Potoni�e 1960

Type species: Droseridites spinosus (Cookson) Potoni�e1960, p. 137–139.

Droseridites senonicus Jardin�e and Magloire, 1965Plate 2, fig. 16

Remark: The present specimens show greater tetrad

diameters than that of the original specimens (12–

19 mm) described by Jardin�e and Magloire (1965).

Maximum tetrad diameter (3 specimens) 19 (21)22 mm.

7. Conclusions

(1) This work provides the first quantitative distri-

butions of Egyptian post-Aptian palynofloral

assemblages, which, combined with semi-quan-

titative data, enables a wider comparison ofassemblages on regional and interregional

scales. The Albian–Cenomanian palynoflora

recovered from the Abu Tunis 1x

1000

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20 A.S. Deaf et al.

TPAL_A_828662.3d (TPAL) 21-08-2013 19:36

borehole shows characteristics of the Albian–

Cenomanian Elaterates Phytogeographic Prov-

ince, whereas the early Santonian palynoflora

is exclusively represented by marine phyto-plankton and completely lacks any terrestrial

palynomorphs representative of the Senonian

Palmae Province.

(2) The qualitative and quantitative distributions

of index palynomorph taxa of the Abu Tunis

1x borehole have enabled the recognition of

four Cretaceous lithostratigraphic units, which

permit new age assignments to sections of theborehole undated by the drilling company.

These four units have been subdivided into five

palynological zones. A barren interzone is

recorded just below the uppermost biozone. An

absence of Turonian–Coniacian palynomorphs

may be related to the unfavourable limestone

and dolostone lithologies. An unconformity

surface located between the barren interval andthe overlying lower Santonian may indicate a

short time cessation in deposition and erosion,

which may be related to the Turonian uplift

and basin inversion of the Faghur area, which

has affected most of the northern basins of

Egypt.

(3) Correlation between the biozones defined for

the Abu Tunis 1x borehole with those ofSchrank and Ibrahim (1995) shows that the

new scheme is applicable at least to Albian–

middle Cenomanian succession in the Western

Desert. However, there are some difficulties

correlating the upper Cenomanian parts of the

successions, which are due to the change in

depositional environments from clastic shallow

marine to carbonate-rich open marine environ-ments in the more northern basins. Further pal-

ynological (preferably independently age

controlled) work on successions in adjacent

and less tectonically affected basins may permit

the development of a complete late Cenoma-

nian–Maastrichtian palynozonal scheme for

the Egyptian northern Western Desert.

(4) A major synchronicity between the North andWest African late Cretaceous biostratigraphic

units is suggested here, in contrast to the argu-

ment made by Schrank and Ibrahim (1995).

The synchronicity is based on occurrence of

foraminifera-calibrated biostratigraphic events

in both the North and West African sequences.

The cause of the synchronicity is strongly sug-

gested to be the continuous mid–late Cenoma-nian anticlockwise north-northeast African

plate movement towards Laurasia as a

response to the breakup of Western

Gondwana. This resulted in North and West

African countries being brought to a similar

palaeosubtropical position and thus under sim-

ilar palaeoclimate, whereby these regionsshould have similar palynofloral compositions

of similar age ranges.

(5) The quantitative vertical distribution of Afro-

pollis jardinus recorded from the Abu Tunis 1x

borehole sediments shows a mid Albian–early

Cenomanian acme event that can be correlated

with similar semi-quantitatively identified

acmes of the species in other wells in the northWestern Desert of Egypt, permitting the recog-

nition of this acme event as an important

regional stratigraphic marker.

Acknowledgements

A.S. Deaf gratefully thanks the Egyptian Government forgenerously funding him a PhD scholarship at University ofSouthampton, UK. The authors wish to thank the EgyptianGeneral Petroleum Corporation for providing well logs andsamples of the Abu Tunis 1x borehole. Thanks are also dueto Mr. Shir Akbari at NOCS, Q7University of Southampton,for his help in the laboratory.

Author biography Q8

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Meshref WM. 1990. Tectonic framework. In: Said R, editor.The Geology of Egypt.Rotterdam: Balkema; p. 113–156.

Morgan R. 1978. Albian to Senonian palynology of Site 364,Angola Basin. Initial Rep Deep Sea. 40:915–951.

M€uller J. 1966. Montane pollen from the Tertiary of NWBorneo. Blumea. 4:231–235.

Norton P. 1967. Rock stratigraphic nomenclature of theWestern Desert. Internal Report. Cairo.Q10

Omran AM, Soliman HA, Mahmoud MS. 1990. Early Creta-ceous palynology of three boreholes from northernWesternDesert (Egypt). Rev Palaeobot Palyno. 66(3–4):293–312.

Penny JHJ. 1986. An early Cretaceous angiosperm pollenassemblages from Egypt. In: Batten DJ, Briggs DEG,editors. Studies in Palaeobotany and Palynology in Hon-our of N. F. Hughes. London: Special Papers in Palaeon-tology; p. 119–132.

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Said R. 1962. Tectonic framework of Egypt. In: Said R, editor.The Geology of Egypt.Amsterdam: Elsevier; p. 28–44.

Salard-Cheboldaeff M. 1990. Intertropical African palynos-tratigraphy from Cretaceous to Quaternary times. J AfrEarth Sci. 11: 1–24.

Schrank E. 1992. Nonmarine Cretaceous correlations inEgypt and northern Sudan; palynological and palaeobo-tanical evidence. Cretaceous Res. 13(4):351–368.

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Schrank E, Mahmoud MS. 1998. Palynology (pollen, sporesand dinoflagellates) and Cretaceous stratigraphy of theDakhla Oasis, central Egypt. J Afr Earth Sci. 26(2):167–193.

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Appendix 1. List of palynological samples from the Abu Tunis 1x borehole showing total recovery of palynomorphs in

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Appendix 2. Alphabetic list of palynomorphs recorded in

the Abu Tunis 1x borehole

Spores

Balmeisporites cf. holodictyus Cookson & Dettmann, 1958 (9).Cicatricosisporites orbiculatus Singh, 1964 (4).Cicatricosisporites sinuosusHunt, 1985 (13).Cicatricosisporites spp. (7).Concavissimisporites spp. (11).Crybelosporites pannuceus (Brenner) Srivastava, 1977 (2).Deltoidospora halliiMiner, 1935 (5).Deltoidospora toralis (Leschik) Lund, 1977 (16).Deltoidospora spp. (1).Gleicheniidites senonicus Ross, 1949 (15).Todisporites minor Couper, 1958 (14).Trilobosporites laevigatus El-Beialy, 1994 (6).Triplanosporites sp. (3).Triporoletes reticulatus (Pocock) Playford, 1971 (12).Verrucosisproites obscurilaesuratus Pocock, 1962 (8).

Gymnosperm pollen

Alisporites cf. grandis (Cookson) Dettmann, 1963 (17).Arucariacites australis Cookson ex Couper, 1953 (24).Balmeiopsis limbatus (Balme) Archangelsky, 1979 (18).Classopollis classoides Pflug, 1953 (19).Classopollis spp. (23).Cycadopites spp. (22).Elaterocolpites castelainii Jardin�e &Magloire, 1965 (25).Elaterosporites acuminatus (Stover) Jardin�e, 1967 (28).Elaterosporites klaszii (Jardin�e &Magloire) Jardin�e, 1967 (21).Elaterosporites protensus (Stover) Jardin�e, 1967 (29).Elaterosporites verrucatus (Jardin�e & Magloire) Jardin�e, 1967(27).Ephedripites irregularisHerngreen, 1973 (33).Ephedripites spp. (20).Eucommidites treodsonii (Erdtman) Potoni�e, 1958 (34).Galeacornea causea Stover, 1963 (31).Gnetaceaepollenites cf. clathratus Stover, 1964 (30).Inaperturopollenites undulatusWeyland & Greifeld, 1953 (32).Sofrepites legouxae Jardin�e, 1967 (26).

Angiosperm pollen

Afropollis jardinusDoyle et al., 1982 (36).Afropollis aff. jardinusDoyle et al., 1982 (66).Afropollis kahramanensis Ibrahim & Schrank 1995 (42).Cretacaeiporites densimurus Schrank & Ibrahim, 1995 (38).Cretacaeiporites mulleriiHerngreen, 1973 (45).Cretacaeiporites polygonalis (Jardin�e & Magloire) Herngreen,1973 (52).Dichastopollenites ghazalatensis Ibrahim, 1996 (55).Foveotricolpites gigantoreticulatus (Jardin�e & Magloire)Schrank, 1987a (57).Papillopollis vancampoaeKedves & Pittau, 1979 (40).Proteacidites cf. africaensis (Jardin�e & Magloire) Schrank &Ibrahim, 1995 (35).Retimonocolpites ghazalii Ibrahim, 2002 (59).Retimonocolpites variplicatus Schrank &Mahmoud, 1998 (41).Retimonocolpites textus (Norris) Singh, 1983 (49).Rousea brenneri Singh, 1983 (50).Rousea delicipollis Srivastava, 1977 (44).Rousea cf. miculipollis Srivastava, 1975 (47).Stellatopollis barghoorniiDoyle in Doyle et al., 1976 (61).Stellatopollis dejaxii Ibrahim, 2002 (64).

Stellatopollis densiornatus (Lima) Ward, 1986 (62).Stellatopollis doylei Ibrahim, 2002 (69).Stellatopollis limai Ibrahim, 2002 (65).Stellatopollis spp. (68).Stephanocolpites sp. (60).Striatopollis cf. trochuensis (Srivastava) Ward, 1986 (63).Tetracolpites sp. (67).Tetraporopollenites sp. (39).Tricolpites cf. crassimurus (Groot & Penny) Singh, 1971 (56).Tricolpites micromunus (Groot & Penny) Singh, 1971 (54).Tricolpites parvus Stanley, 1965 (51).Tricolpites sagaxNorris, 1967 (58).Tricolpites vulgaris (Pierce) Srivastava, 1969 (48).Tricolpites spp. (37).Tricolporopollenites sp. (43).Triporites spp. (46).Triporopollenites spp. (53).

Pollen tetrads

Cretacaeiporites densimurus Schrank & Ibrahim, 1995 (70).Droseridites baculites Ibrahim, 1996 (71).Droseridites senonicus Jardin�e & Magloire, 1965 (72).

Freshwater algae

Botryococcus sp. (75).Chomotriletes minor (Kedves) Pocock, 1970 (76).Fungal fruiting body (74).Ovoidites parvus (Cookson &Dettmann) Nakoman, 1966 (73).

Dinoflagellate cysts

Canningia senonica Clarke & Verdier, 1967 (79).Cannosphaeropsis utinensisWetzel, 1933 (96).Chatangiella madura Lentin & Williams, 1976 (81).Chlamydophorella discreta Clarke & Verdier, 1967 (91).Circulodinim distinctum (Deflandre & Cookson) Jansonius,1986 (100).Coronifera albertiiMillioud, 1969 (101).Coronifera oceanica Cookson & Eisenak, 1958 (97).Coronifera tubulosa Cookson & Eisenak, 1974 (104).Coronifera spp. (113).Cribroperidinium edwardsii (Cookson & Eisenack) Davey,1969 (108).Cribroperidenium sp. (110).Dinogymnium denticulatum (Alberti) Evitt et al., 1967 (89).Dinogymnium spp. (82).Downiesphaeridium sp. (92).Eucladinium gambangense (Cookson & Eisenack) Stover &Evitt, 1978 (93).Exochosphaeridium bifidum (Clarke & Verdier) Clarke et al.,1968 (80).Florentinia berran Below, 1982 (98).Florentinia laciniataDavey & Verdier, 1973 (103).Florentinia mantellii (Davey & Williams) Davey & Verdier,1973 (99).Florentinia spp. (94).Isabelidinium acuminatum (Cookson & Eisenack) Stover &Evitt, 1978 (90).Litosphaeridium siphoniphorum (Cookson & Eisenack) Davey&Williams, 1966 (86).Odontochitina operculata (Wetzel) Deflandre & Cookson,1955 (78).

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Odontochitina costata Alberti, 1961 (83).Odontochitina porifera Cookson, 1956 (84).Oligosphaeridium albertense (Pocock) Davey & Williams,1969 (105).Oligosphaeridium complex (White) Davey & Williams, 1966(106).Oligosphaeridium poculum Jain, 1977 (107).Pseudoceratium anaphrissum (Sarjeant) Bint, 1986 (111).Pseudoceratium securigerum (Davey & Verdier) Bint, 1986(112).Pterodinium sp. (102).Senegalinium aenigmaticum (Boltenhagen) Lentin & Wil-liams, 1981 (85).Spinifereites spp. (77).Subtilisphaera senegalensis Jain & Millepied, 1973 (109).Surculosphaeridium cf. longifurcatum (Firtion) Davey et al.,1966 (88).Trichodinium castaneaDeflandre, 1935 (87).Xiphophoridium alatum (Cookson & Eisenack) Sarjeant, 1966(95).

Acritarchs

Baltisphaeridium spp. (114).Micrhystridium spp. (115).

Microforaminiferal test linings (116)

Albian angiosperm pollen grains

1–5. Afropollis jardinus Doyle et al. 1982, 1, sample 89/A,R13; 2, sample 110/A, L46/3; 3, sample 89/B, Q22; 4, sample93/A, N19/3; 5, sample 110/A, L46/3, (36).

Mid Albian elaterate gymnosperm pollen grains

6, 8. Elaterosporites verrucatus (Jardin�e & Magloire) Jardin�e1967, sample 88/B, M31/3, sample 93/A, J23/3, (27).

7, 14. Elaterosporites acuminatus (Stover) Jardin�e 1967,sample 93/A, G45/4, sample 91/A, H32, (28).

9. Elaterosporites protensus (Stover) Jardin�e 1967, sample90/A, U16, (29).

10. Elaterosporites klaszii (Jardin�e & Magloire) Jardin�e1967, sample 97/A, G13/4, (21).

Late Albian elaterate gymnosperm pollen grains

13, 15–17. Sofrepites legouxae Jardin�e 1967, 13, sample 93/A,M32/3; 15, sample 86/B, Q30; 16, sample 93/A, T23/2; 17,sample 95/A, H17/3, (26).

Late Albian–Mid Cenomanian pollen grains

11. Cretacaeiporites mullerii Herngreen 1973, sample 103/A,U25, (45).

12. Cretacaeiporites polygonalis (Jardin�e & Magloire)Herngreen 1973, 94/A, K32/3, (52).

18. Ephedripites irregularis Herngreen 1973, sample 77/B,C19/2, (33).

19. Gnetaceaepollenites cf. clathratus Stover 1964, sample83/A, K44/1, (30).

20. Galeacornea causea Stover 1963, sample 84/A, K46,(31).

21. Afropollis kahramanensis Ibrahim & Schrank 1995,sample 88/A, Q38/1, (42).

22. Foveotricolpites gigantoreticulatus (Jardin�e &Magloire) Schrank 1987, sample 92/A, L13/3, (57).

Albian–Cenomanian spore and pollen grains

1. Tetracolpites sp., sample 77/B, G28/2, (67).2. Tricolpites sagaxNorris 1967, sample 92/A, G29, (58).3. Striatopollis cf. trochuensis (Srivastava) Ward 1986,

sample 86/A, Q42/4, (63).4. Stephanocolpites sp., sample 91/A, Q23/3, (60).5. Tricolpites parvus Stanley 1965, sample 94/A, M36,

(51).6. Tricolpites cf. crassimurus (Groot & Penny) Singh

1971, sample 85/A, J20/4, (56).7. Rousea delicipollis Srivastava 1977, sample 71/A, D34/

2, (44).8. Tricolpites micromunus (Groot & Penny) Singh 1971,

sample 75/A, Q43/3, (54).9. Triporopollenites sp., sample 94/A, S45, (53).10. Rousea cf. miculipollis Srivastava 1975, sample 97/A,

P38/2, (47).11. Proteacidites cf. africaensis (Jardin�e & Magloire)

Schrank & Ibrahim 1995, sample 85/A, O42, (35).12. Tricolporopollenites sp., sample 93/B, W14/3, (43).15. Droseridites baculites Ibrahim 1996, sample 92/A,

Q40/4, (71).16. Droseridites senonicus Jardin�e & Magloire 1965, sam-

ple 77/B, P27/4, (72).17. Crybelosporites pannuceus (Brenner) Srivastava 1977,

sample 91/A, L44, (2).

Cretaceous spore and pollen grains

13, 14, 19–22. Retimonocolpites textus (Norris) Singh 1983, 13and 14, sample 94/A, N28/3; 19 and 20, sample 89/A, K40/1;21, sample 94/A, G31/4; 22, sample 94/A, G31/4, (49).

18. Dichastopollenites ghazalatensis Ibrahim 1996, sample88/B, X37/2, (55).

24. Retimonocolpites variplicatus Schrank & Mahmoud1998, sample 90/A, Q30, (41).

Fresh water algae

23. Botryococcus sp., sample 99/B, R42/4, (75).25. Fungal fruiting body, sample 109/B, N34/4, (74).

Santonian dinoflagellate cyst

1. Canningia senonica Clarke & Verdier 1967, sample 131/B,L22, (79).

Post-Turonian dinoflagellate cysts

2, 6, 15. Dinogymnium denticulatum (Alberti) Evitt et al.1967, sample 130/A, Q36, sample 130/A, P42, sample 130/B,N29/4, (89).

3, 7. Eucladinium gambangense (Cookson & Eisenack)Stover & Evitt 1978, sample 130/A, K28, sample 130/B, Q42/4,(93).

4, 13. Isabelidinium acuminatum (Cookson & Eisenack)Stover & Evitt 1978, sample 131/A, U27/2, sample 131/B,M34, (90).

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9. Dinogymnium sp., sample 130/B, S45/1, (82).10. Dinogymnium sp., sample 131/B, U26/3, (82).12. Chatangiella madura Lentin & Williams 1976 sample

134/A, U26, (81).14. Exochosphaeridium bifidum (Clarke & Verdier) Clarke

et al. 1968 sample 133/A, C11/3, (80).17. Cannosphaeropsis utinensis Wetzel 1933, sample 117/

A, S33/3, (96).19. Odontochitina porifera Cookson 1956, sample 133/A,

L39/1, (84).

Cretaceous dinoflagellate cysts

5. Trichodinium castanea Deflandre 1935, sample 116/A, L12/4, (87).

8. Xiphophoridium alatum (Cookson & Eisenack) Sarjeant1966, sample 103/B, E43/4, (95).

11, 18. Senegalinium aenigmaticum (Boltenhagen) Lentin& Williams 1981, sample 110/A, G13/3, sample 109/A, S31/1,(85).

16. Odontochitina costata Alberti 1961, sample 134/A,V38/2, (83).

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