Palynology and Palaeoenvironment of the Quseir Formation (Campanian) from Central Egypt Magdy S

Journal of African Earth Sciences 36 (2003) 135–148 www.elsevier.com/locate/jafrearsci

Palynology and palaeoenvironment of the Quseir Formation (Campanian) from central Egypt Magdy S. Mahmoud Department of Geology, Faculty of Science, Assiut University, Assiut 71516, Egypt Received 19June 2002; accepted 9May 2003

Abstract The palynofloras of the basal part of the Quseir Formation in the Bulaq area, central Egypt, are overwhelmingly of terrestrial origin. They are dominated by angiosperms (mainly Foveotricolpites and Arecipites). Pteridophytic spores are abundant, amongest which the Deltoidospora/Cyathidites association and Gabonisporis vigourouxii are the most frequent. Aquatic plants (e.g. Ariadna- esporites spores) and freshwater algae (e.g. Ovoidites and Pediastrum) occur in appreciable amounts. The association is indicative of a fluvio-lacustrine environment characterized by widespread moist and aquatic habitats under a warm-humid (tropical) palaeoclimate. An angiosperm-based dating as Campanian (most probably Early Campanian) is suggested. Proteacidites sp. 3 Lawal and Moullade and Syncolporites schrankii Awad are the most significant angiosperms, which are not known to range before the Campanian in the ‘‘Senonian Palmae Province’’ areas. The Bulaq assemblages bear a close relationship with the Palmae palynofloras of North Africa, but differ significantly from those of West Africa. Ó 2003 Elsevier Ltd. All rights reserved.

Keywords: Terrestrial palynology; Palaeoenvironment; Quseir Formation; Campanian; Egypt

1. Introduction The stratigraphic position of the basal beds of this unit is significant because they rest unconformably over The general Kharga-Dakhla field area investigated Turonian (Taref Formation) rocks. The Coniacian– here was the subject of several palynological investiga- Santonian interval is represented, however, by a major tions since the early sixties (Helal, 1965). In most recent hiatus in Central Egypt (Klitzsch and Hermina, 1989, publications (e.g. Schrank and Mahmoud, 1998, 2000, p. 83). In the present paper, well-preserved terrestrial 2002), well-preserved palynomorphs were extracted palynomorphs of Campanian age are described from the from the clay/shale beds intercalated with the Nubia- shale of the Quseir Formation. The material comes type siliciclastics in the Dakhla-Kharga areas. These from the basal part of the formation in the subsurface rocks were previously regarded as ‘‘unfossiliferous’’; succession of the Bulaq area (Fig. 2). Due to lack of mar- their terrestrial to marginal marine origin prevents in- ker angiosperm taxa, Mahmoud (1998) dated a paly- dependent faunal dating. Although the palynostrati- nomorph association from Kharga as ‘‘Campanian’’ but graphic record of these rocks is fragmentary, new this was later interpreted by Schrank and Mahmoud stratigraphic and taxonomic information has been de- (2000) as Late Cenomanian (equivalent to the Maghrabi rived from the study of the fossil palynomorphs. But Formation). The present palynomorph assemblage of- establishment of a complete palynological record has fers direct and strong biostratigraphic evidence as to the been hindered by the siliciclastic nature of these rocks age and palaeoenvironment of the lower Quseir For- (predominantly sandstones and siltstones) which are not mation. In addition to biostratigraphic and palaeo- suitable for palynomorph recovery. However, palyno- ecologic investigations, which are the main goals, the floras were not described yet from the extensive Quseir present work aims also to compare the Bulaq palyno- (shale) rocks (surface and subsurface) in central Egypt. floras with their contemporary counterparts from the Palmae province. Because the palynomorph assemblage E-mail addresses: [email protected], [email protected] is diverse and well preserved it will contribute to the (M.S. Mahmoud). Palmae palaeofloristic characteristics.

0899-5362/03/$ - see front matter Ó 2003 Elsevier Ltd. All rights reserved. doi:10.1016/S0899-5362(03)00047-2 136 M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 2. Geographic and geological setting Formation (see Klitzsch and Hermina, 1989, p. 126, 127). The Quseir Formation contains locally remains of The Quseir Formation (Youssef, 1957) in the sur- gastropods, lungfishes (Ceratodus tuberculatus and C. roundings of Kharga (Fig. 1), is the uppermost unit humei), sawfishes (Schizorhiza stromeri) and mosasaur- of the former ‘‘Nubia Sandstone’’ succession (Klitzsch ids and turtles (Podocnemis sp.), which might indicate et al., 1979), and consists of an alternation of claystone, Early to Late Campanian age. These fossil associations siltstone and sandstone beds. It was also named as the in the Quseir Formation, however, did not provide a Variegated Shales (Said, 1962) and as the Mut Forma- definite age. Previous palynological work in the Kharga tion (Barthel and Herrmann-Degen, 1981). Coeval to Oasis area (Soliman, 1977; Mohsen, 1992) gave incon- the formation is the Wadi Hennis Formation (Dominik, clusive dating for the basal beds from the Bulaq area 1985) at the Farafra Oasis area to the north of Kharga. because the assemblages lack marker angiosperms. The Quseir Formation overlies the Taref Formation These alleged ages were given, for the Bulaq-15 (depth (Turonian) with a conspicuous unconformity and is 206–212 m) and Bulaq-12 (depth 202–208 m) cores, as slightly unconformably overlain by the well dated Upper Cretaceous, probably Cenomanian and for the (Campanian to Early Maastrichtian) marine Duwi Bulaq-14 core (depth 212–214 m) as Albian (Soliman, 1977). Mohsen (1992) gave a rather different age for the Bulaq-15 core (depth 206–224 m) as Coniacian and suggested a nonmarine palaeoenvironment. For additional information regarding the stratigraphy of the Nubia strata in general, and the Quseir Formation in particular, refer to Klitzsch et al. (1979), Klitzsch and Hermina (1989), Klitzsch and Squyres (1990) and Hermina (1990). In the surroundings of Kharga Oasis the base of the Quseir Formation exhibits an inferred marginal marine (tidal flat) palaeoenvironment which grades upward to shallow marine (see Klitzsch and Hermina, 1989). The basal shale and sandstones of the formation in these areas provides evidence of a terrestrial and brackish palaeoenvironment (Hermina, 1990). Awad and Ghob- rial (1965) and Hendriks et al. (1984) previously inferred limnic conditions in the basal sediments of the formation, where freshwater gastropods, abundant plant debris and other remains of reptiles and dinosaurs are found.

3. Materials and methods

Core samples from two borehole sections (Bulaq-12 and Bulaq-15) from the Bulaq area, Kharga Oasis are investigated palynologically (Figs. 1 and 2). Because these sections are siliciclastics of Nubia type, the ma- jority of the processed samples proved to be barren of palynomorphs. Five samples contain palynomorphs suitable for the present work. Samples were digested by acids (35% HCl to dissolve the carbonates and 40% HF to remove the silicates). No centrifugation, heavy liquid separation and ultrasonic procedures were made. In addition, the residues were not subjected to oxidation or Fig. 1. Geological map of the Kharga Oasis (after Klitzsch et al., alkali treatments. The acid-digested residues were sieved 1987) showing locations of the two boreholes (Bu12 ¼ Bulaq-12; using 10 lm polyamide nylon sieves. Permanent slides Bu-15 ¼ Bulaq-15). From top to bottom stratigraphic units are Q ¼ were prepared using glycerine jelly as a mounting me- Quaternary; T ¼ Tertiary; Kud ¼ Dakhla Formation; Kuw ¼ Duwi Formation; Kuq ¼ Quseir Formation, Kut ¼ Taref Formation; Kum ¼ dium. For each sample ﬁve permanent slides were pre- Maghrabi Formation; Kls ¼ Sabaya Formation. For ages of these pared. A count of 200 grains was achieved in each units see Klitzsch and Hermina (1989). sample for the purpose of making a semi-quantitative M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 137

Fig. 2. Lithological logs of two borehole sections (logs modified after unpublished charts by the Hydrology Section of the Egyptian General Desert Development Organization, Management of the Groundwater at Kharga) with positions of palynomorph-productive core samples. estimate of palynomorphs. Additional slides were in- colpites). Other tricolpates (Tricolpites) and monocol- vestigated qualitatively. The washed residues were pates (Stellatopollis) occur but are of lower diversities. stored with distilled water in small glass bottles. Slides Triporates (Proteacidites) and syncolporates (Syncolpor- and residues are stored with the palynological collec- ites) occur sporadically. Tricolporates such as Reti- tions of the Geological Museum of the University of tricolporites are minor constituents. The spores are Assiut. represented mainly by a diverse group of ferns of the Deltoidospora/Cyathidites association, which include several species of negligible stratigraphic value. Other 4. Results ferns such as Biretisporites, Concavisporites, Triplanosp- orites and others occur, but are not as abundant and 4.1. Palynoflora diverse as Deltoidospora/Cyathidites representatives. Bryophyte spores (Zlivisporis) are rare, but in sample 1 Qualitative data. The palynomorphs of the Bulaq area they are noticeably abundant. Gymnosperms, freshwater are well preserved, in which the angiosperms and the algae and fungal spores, although abundant, are only spores are the main and most diverse constituents. The represented by a few genera. Gymnosperms encompass angiosperms are represented essentially by several species representatives from the araucariaceans, ephedroids and of Foveotricolpites and by monocolpates (Arecipites, monosulcates (Monosulcites) in all samples. A single grain Foveomonocolpites, Monocolpopollenites and Retimono- of Cingulatipollenites occurs in sample 2. Unidentifiable 138 M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 forms of freshwater algae such as rounded (presinophyte- West Africa and northern South America. In general, like) and other rectangular (Fig. 7S) types dominate over the Late Cretaceous (Turonian-Senonian) assemblages others. Two species of Ovoidites, in addition to some exhibit great similarities in their palynofloral charac- sporadic presence of Botryococcus, Chomotriletes and teristics in the local palynofloras. Significant biostrati- Pediastrum are other freshwater algae that are found in graphic taxa are, in most cases, rare. They are rarely the Bulaq associations. Fungal spores and fungal fruiting seen from palynological specimens. In the Lower Se- bodies are common constituents of the palynoflora. nonian, an upward increase in the frequency of Palmae- Quantitative data. In a broader sense, the Bulaq paly- type monocolpate pollen occurred. A local (Turonian to nofloras as seen from their semi-quantitative distribution Santonian) association (Ibrahim and Abdel-Kireem, (Table 1) and diagramatic presentation (Fig. 3) are gene- 1997; Farafra Oasis, Egypt) is a good example: the as- rally similar in the five studied samples. However, some sociation was dated on the basis of few taxa in which minor differences were observed; maximum abundance of tetrads of Droseridites senonicus were the most impor- the araucariacean and ephedroid pollen occurs in sample tant elements. The present association is typical of the 3. Also, the least angiosperms are seen in sample 1 Senonian Palmae Province (Herngreen and Chlonova, (26.5%). The palynoflora in general is dominated by the 1981; Herngreen et al., 1996). Palynofloras of this angiosperms (up to 51.6% of total palynomorphs, sample province encompass several contemporary palyno- 4). The foveolate tricolpates (Foveotricolpites) and the morphs that are found in rocks of Coniacian to Maas- monocolpates (mainly Arecipites) are the most abundant. trichtian ages (e.g. Herngreen et al., 1996). This is clearly Proteacidites sp. 3, questionable Retidiporites, Syncolpo- seenfromtheoccurrenceofPalmae(e.g.Mono- rites schrankii and S. subtilis are rare, although otherwise colpopollenites) as well as other proteacean (e.g. Proteaci- significant from a stratigraphic point of view. Spores, dites) and syncolporate (e.g. Syncolporites) angiosperms. gymnosperms as well as freshwater algae are well repre- Pre-Campanian (e.g. Turonian) assemblages normally sented but are not as abundant as angiosperms. Gabo- contain abundant tricolp(or)ates with others such as nisporis (up to 17.3%, sample 1) and pteridophytic spores members of the Foveotricolpites giganteus/gigantoreti- (mainly Deltoidospora/Cyathidites, up to 14.9%, sample 1) culatus group. dominate the spores. Bryophyte spores occur sporadically In North Africa (Egypt, Sudan and Somalia) several in all samples but in sample 1 they reach up to 12.9% of peculiar Campanian-Maastrichtian angiosperm taxa total palynomorphs. Gymnosperms (mainly araucaria- such as Buttinia, Echitriporites, Gemmatricolpites, Peri- cean and ephedroid pollen) reach up to 25.8% (sample 3). retisyncolpites (an endemic African pollen) and Spinizo- Ephedroids, however, are rare constituents in Bulaq-12 nocolpites occur (Schrank, 1984, 1987a,b, 1994a; El palynoflora (1.2%, sample no. 1), but in Bulaq-15 they Beialy, 1995). These taxa are absent in the present as- exhibit greater percentages (up to 16.5%, sample 3). semblage. The basal Quseir palynoflora might therefore Freshwater algae (up to 12.6%, sample 2) are also im- predate these angiosperms. On the other hand, the West portant members of the palynomorph content (see Fig. 3). African assemblages contain other significant angiosperms that are not known in North Africa, such as 4.2. Palynofacies Hexacolpites, Hexaporotricolpites, Retistephanocolpites and many others (e.g. Salard-Cheboldaeff, 1990). The palynofacies is dominated by large wood (brown Influence of other palynofloral provinces such as the and black) phytoclasts and, less commonly, cuticle sheets. northern Late Turonian and Senonian Aquilapollenites Cuticles possess polygonal cell outlines, sometimes with Provinces (Herngreen and Chlonova, 1981; Herngreen irregular outlines. Both wood phytoclasts and cuticles are et al., 1996) can be seen from the occurrence of the well preserved. Structured wood phytoclasts and trac- Proteacidites pollen and Syncolporites subtilis. S. sub- heids are rare. Amorphous organic matter and other tilis, however, is a member of the Callistopollenites membranous structures are found rarely. Degraded or- group sensu Frederiksen (1987). It is also compared to ganic remains of smaller sizes are very rare. Palyno- Cranwellia striata, another member of this group. morphs (miospores, freshwater algae and some fungal hyphens) are not as common as organic debris. These 5.2. Palynological age of the basal Quseir Formation characteristics are, more or less, found in all samples. Most of the present pollen and spore species have been recorded from Upper Cretaceous rocks in West, 5. Discussion Central and North Africa as well as in other parts of the Senonian Palmae Province. These include different spe- 5.1. The Bulaq palynoflora in a local and regional context cies of angiosperms such as Foveotricolpites, Arecipites, Liliacidites and spores such as Gabonisporis and Zli- Several Bulaq pollen and spore species are known visporis blanensis. The base of the Campanian-Early from the Turonian and Senonian rocks of North and Maastrichtian Zone V (Lawal and Moullade, 1986) M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 139

Table 1 Semi-quantitative distribution of palynomorphs encountered in the basal part of the Quseir Formation, Bulaq area Core no. 12345 Spores Acanthotriletes sp. Ariadnaesporites sp. Fig. 5/E djjdd Biretisporites potoniaei Delcourt and Sprumont d Camarozonosporites sp. Fig. 5/G Cavate spores (Balmeisporites-like) Fig. 5/I dd Cicatricosisporites sp. Concavisporites sp. Fig. 5/K Concavissimisporites sp. Deltoidospora/Cyathidites Group Fig. 5/J jjjjj Foveotriletes margaritae (Van der Hammen) d Gabonisporis sp. cf. G. labyrinthus Srivastava, Fig. 5/M dd Gabonisporis vigourouxii Boltenhagen, 1967 Fig. 5/A N jdjj cf. Lycopodiumsporites sp. Fig. 5/B d Matonisporites equiexinus Couper dd Todisporites major Couper ddd Triplanosporites sp. ddd d Verrucosisporites sp. Fig. 5/N d Zlivisporis blanensis Pacltova, 1961 Fig. 5/C, D N Freshwater algae aﬀ. Celyphus sp. Fig. 7/S d Botryococcus sp. Fig. 7/W Chomotriletes minor (Kedves) Fig. 7/M Ovoidites ligneolus Potonie ex Krutzsch, Fig. 7/V Ovoidites parvus (Cookson and Dettmann) Fig. 7/U dd Pediastrum sp. d Prasinophyte-like freshwater algae (not ﬁgured) jjdjj Fungal spores Pluricellaesporites sp. Fig. 7/T d Rhyzophagites sp. Fig. 7/R Other fungal spores (Fig. 5/L) and fruiting bodies jdddj Indet. palynomorphs Palynomorph type A Fig. 5/F Palynomorph type B Fig. 5/H d Palynomorph type C Fig. 6/E d Gymnosperm pollen Araucariacean pollen Fig. 5/Q ddjdd Cingulatipollenites sp. Ephedroid pollen Fig. 5/O, P djN jj Monosulcites sp. Fig. 5/R d ddd Angiosperm pollen Arecipites sp. Fig. 6/I jdd d cf.Cretacaeiporites polygonalis (Jardine and Magloire) Fig. 6/A Dichastopollenites? sp. sensu Ibrahim and Abdel-Kireem, 1997 Fig. 5/T Foveomonocolpites sp. sensu Schrank, 1994; Fig. 7/B Foveomonocolpites sp.1 Fig. 6/P Foveomonocolpites sp.2 Fig. 6/B dddd Foveotricolp. gigantoreticulatus (Jardine and Magloire) Fig. 6/Kand Fig. 7/F d NNNj F. irregularis Herngreen, 1975 Fig. 7/K F. tienabaensis (Jardine and Magloire) Fig. 7/C d dd Foveotricolpites sp. cf. F. giganteus (Jardine and Magloire) Fig. 7/D, G djdjj Foveotricolpites sp. dd Liliacidites spp. Fig. 6/J, Q dd Longapertites sp. Fig. 7A d Momipites sp. sensu Schrank and Ibrahim, 1995 Fig. 7/O Monocolpopollenites sphaeroidites Jardine and Magloire, 1965 Fig. 6/L dddd Monocolpopollenites sp. dd Pollen tetrads (tetrahedral type as in Droseridites) Fig. 5/S, U d Proteacidites sp. Fig. 6/D (continued on next page) 140 M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148

Table 1 (continued) Core no. 1 2 3 4 5 Proteacidites sp. sensu Schrank, 1994; Fig. 7/I Proteacidites sp. 3 Lawal and Moullade, 1986 Fig. 7/J d cf. Retidiporites magdalen. V. d. Hammen and Garcia, Fig. 6/F d Retidiporites? sp. Fig. 6/M dd Retimonocolpites Group sensu Herngreen, 1975 Fig. 6/C, O dd Retitricolporites fekelensis Meeon, Fig. 7/E cf. Stellatopollis sp. Fig. 6/H dd Syncolporites schrankii Awad, 1994 Fig. 6/G S. subtilis Boltenhagen, 1976 Fig. 6/N Tricolpites maximus (Singh) Fig. 7/L T. microreticulatus Belsky, Boltenhagen and Potoniee, Fig. 7/H dd d Tricolpites sp. S.CI. 278 Jardine and Magloire, 1965 Fig. 7/Q Tricolpites spp. Fig. 7/N dd Tricolporate pollen (small-sized, not ﬁgured) d For full reference to spores and pollen taxa, as for the text as well, see Ravn (1998). Single specimen; d Present (2–10 specimens); j Common (>10 specimens); N Abundant (>30 specimens).

Fig. 3. Percentages of selected palynomorphs and palynomorph groups in samples from the Bulaq boreholes. boundary was considered to be equivalent with the last and Moullade, 1986; Periretisyncolpites, Spinizonocol- occurrence of D. senonicus. This zone contains Retidipo- pites, Syndemicolpites and others (e.g. Schrank, 1987a,b; rites magdalenensis, Monocolpopollenites sphaeroidites El Beialy, 1995) are absent. A Bulaq core (Bu-15, depth and some proteaceans. Among the spores, Zlivisporis 206–224 m), without D. senonicus, was allegedly dated as blanensis attains its maximum occurrence. In a dinocyst- Coniacian by Mohsen (1992) although he identified, dated palynoflora from Coote^ dÕIvoire (Tea-Yassi et al., among others, Proteacidites sigalii, which is typical post 1999) ephedroids and araucariaceans are restricted to Coniacian–Santonian pollen. Although P. sigalii, a rare the Early Campanian, together with proteacean pollen. species, is not seen in the present material, MohsenÕs Zlivisporis (sometimes as Triporoletes) blanensis and finding supports the present Campanian dating. This Gabonisporis vigourouxii range to the Lower Maas- dating is, however, consistent with, and strongly sup- trichtian. Based on that and on the synopsis of the ports, previous age determinations by faunas (cf. ranges of several selected sporomorph taxa (Table 2) a Klitzsch and Hermina, 1989) which indicate also a Campanian age is suggested for the basal part of the Campanian age. Quseir Formation. As seen from this table, Proteacidites sp. 3 Lawal and Moullade and Syncolporites schrankii 5.3. Palaeoecological and palaeoclimatological interpre- Awad did not enter the record before the Early Camp- tations anian. An Early Campanian age is most probable because typical Campanian (and Maastrichtian as well) Implications from palynoflora. The palynomorphs pollen such as Echitriporites, Longapertites sp. 3 Lawal from the basal part of the Quseir Formation, Bulaq M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 141

Table 2 African ranges of selected spores and pollen encountered in Bulaq-12 and Bulaq-15 boreholes Age Spores and pollen 01Foveo- 02Foveo- 03Foveo- 04Gabo- 05Mono- 06Prote- 07Retidipo- 08Syn- 09Syn- 10Zlivisp- tricolpites tricolpites tricolpites nisporis colpopolle- acidites rites colporites colporites oris blan- giganteus gigantore- tienabaen- vigourouxi nites sp.3 magdalen- schrankii subtilis ensis ticulatus sis sphaeroi- ensis dites Maastrichtian j jjjj jjj j jjjj jjj

Campanian jj jjjj jjj jj jjjj jjj

Santonian jjjjj jj jjjjj jj

Coniacian jjjjj jj jjjjj jj

Turonian jjj j j jj j j

Sources of ranges are as follows: Aboul Ela (1978) (8); Awad (1994) (5, 7, 8, 10); Azeema and Boltenhagen (1974) (1); Boltenhagen (1967) (4), 1975 (5), 1976 (5, 9), 1980 (1, 5); El-Beialy (1994) (1); Herngreen and Chlonova (1981) (4, 7, 10); Ibrahim et al. (1995) (1, 2); Ibrahim and Abdel-Kireem (1997) (2, 3, 10); Jan Du Chêene (1977) (7); Jan Du Chêene et al. (1978) (1, 4); Jardine and Magloire (1965) (1, 2, 3, 5, 9); Kaska (1989) (7); Kuyl et al. (1955) (9); Lawal (1982) (1, 2); Lawal and Moullade (1986) (2, 4, 5, 6, 9, 10); Morgan (1978) (1); PetrosÕyants and Trofimov (1971) (1); Regali et al. (1974) (1); Salami (1985) (7); Salami (1984, 1990) (10); Schrank (1984) (7); 1987a (1, 2, 3); 1987b (8 as cf. Syndemicolpites typicus); 1994b (7); Schrank and Ibrahim (1995) (1, 2); Sultan (1985) (1); Tea-Yassi et al. (1999) (4, 10). area, are exclusively derived from terrestrial land plants; composition of palynoflora, the botanical affinities of no unequivocal brackish or normal marine palyno- some spores and pollen entities (Table 3) are also used to morphs (e.g. dinoflagellate cysts and microforaminiferal support interpretations. Likewise, the qualitative de- linings) have been found. The association includes ele- scription of the organic remains (i.e. palynofacies), de- ments that are tolerant of varying terrestrial sites and termined through transmitted-light microscopy, offers sensitive to different palaeoclimates (e.g. different an- additional evidence. giosperm families, ferns, water ferns and xerophytes). Spores of Gabonisporis are similar to Crybelospo- Therefore palaeoenvironmental interpretations are based rites, and the latter is related to the Marsiliaceae almost entirely on the palaeoecological preferences of (Dettmann, 1963; Collinson, 1991). Schrank and Mah- terrestrial palynomorphs. Beside the quantitative bulk moud (1998) noted that Crybelosporites is comparable

Table 3 Botanical affinities of some selected spores and pollen taxa that are significant in their bulk composition, basal part of the Quseir Formation, Bulaq area Spores Ariadnaesporites Salviniaceae (Hall, 1975) Biretisporites, Concavisporites, Concavissimisp- Several fern families such as Matoniaceae, Cyatheaceae and Dicksoniaceae (e.g. Van Erve orites, Deltoidospora/Cyathidites, Foveotriletes, and Mohr, 1988) Matonisporites, Todisporites, Verrucosisporites Gabonisporis Possibly similar to Crybelosporite, a marsilean water fern (Dettmann, 1963; Collinson, 1991). Crybelosporites is comparable with hydropteridacean spores such as Perotrilites striatus and Dettmannaesporites (Schrank and Mahmoud, 1998) Triplanosporites Gleichenoid? (Pflug in Thomson and Pflug, 1953) Zlivisporis Similar to the Hepaticae Oxymitra paleacea (Pacltova, 1961) Angiosperms Arecipites Palmae (Nichols et al., 1973) Foveomonocolpites Possibly Palmae (by analogy with Monocolpopollenites) Foveotricolpites Unknown Liliacidites Liliaceae Longapertites Probably Palmae (Germeraad et al., 1968; Adegoke et al., 1978) Monocolpopollenites Palmae or Cycadales (Nichols et al., 1973) Tricolpites Hamamelidaceae (Singh, 1971; Srivastava, 1977) or Haloragaceae (Schrank, 1994a) 142 M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 with hydropteridacean spores such as Perotrilites stria- ages were observed in the sediments of the Bulaq-12 tus Cookson and Dettmann or Dettmannaesporites borehole (sample 1, 202–208 m). Kedves. Ariadnaesporites (Salviniaceae), Gabonisporis, However, the Bulaq palynofloras are well preserved Zlivisporis blanensis, which are the products of aquatic and highly diverse, and this implies that the vegetation plants, together with other water ferns suggest, there- was not distant from the depositional site. This also fore, that freshwater habitats were extant during depo- means that a pronounced dryness, which leads to poorly sition of the basal parts of the Quseir Formation, at least diverse palynoflora, was not pertinent. During the at the Bulaq area. Campanian-Maastrichtian, the near-equator position of Abundant pteridophytes (mainly several fern species Africa and northern South America (Smith et al., 1982), of the Deltoidospora/Cyathidites association) in the where the Palmae Province is located, supports a warm, studied samples suggest growing of widespread herba- tropical to subtropical climate for the Kharga region. ceous hygrophilous plants on moist biotopes. Abundant The vegetation realm correlates essentially with one of araucariacean pollen in sample 3 may suggest the oc- lowland, moist to aquatic biotopes where palms, her- currence of araucariacean forests during periods of high baceous fern plants, water ferns, hepatics and other moisture (e.g. Schrank, 1994b). This is supported by the freshwater algae could flourish, sometimes with abun- presence of the most abundant Salvinialean water fern dant conifer (araucariacean) vegetation during the high Ariadnaesporites in the same sample. Araucariaceans, moisture episodes in the relatively dry hinterlands. however, may grow on relatively dry lands (cf. Schrank The absence of Nypa-like mangrove pollen (e.g. Spin- and Mahmoud, 1998). izonocolpites) and marine dinocysts supports a conti- Deposition in such limnic conditions, and moist nental setting of the study area during the Campanian. biotopes for the basal Quseir Formation in the Bulaq Spinizonocolpites was developed during Campanian- area, are supported by the occurrence of freshwater al- Maastrichtian transgressive periods; it is a common gae (with Ovoidites) and hepatic spores. These plant mangrove palm found in coastal basins of North Africa communities thrive today in or near water bodies (see (e.g. Schrank, 1994b). This might imply that the Late Schrank and Mahmoud, 1998). Dry habitats and/or Cretaceous sea reached central Egypt (where the Bulaq seasonal dry periods might have occurred during depo- area is located) relatively later. Otherwise, the previous sition and reached a remarkable development, which recognition of brackish environment, as recorded from correspond to the highest ephedroids (see Table 1). In the base of the Quseir Formation in the Kharga area other words, these might have been xerophytes, which (Hermina, 1990), could be interpreted as an arm of the were adapted to moist conditions. Present day ephe- Tethys which transgressed southward. droids live in semi-arid and dry habitats (e.g. Schrank Implications from palynofacies. Palynofacies analysis and Nesterova, 1993). The lowest Ephedripites percent- can also be used to interpret the palaeoenvironment of

Fig. 4. (a) Well-preserved cuticle phytoclast of irregular cell outlines, X 400. (b) Well-preserved biostructured phytoclast, X 400. (c) Terrestrial palynofacies dominated by wood phytoclasts (brown and black), with rare palynomorphs, X 100. (d) Palynofacies dominated by wood and cuticular phytoclasts, preservation good, with other membranous structures, X 170. M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 143

Fig. 5. For Figs. 5–7 magniﬁcations are X 500 unless otherwise indicated. (A) Gabonisporis vigourouxii, sample 3. (B) cf. Lycopodiumsporites sp., sample 1, X 1200. (C and D) Zlivisporis blanensis ((C) sample 4, (D) sample 1). (E) Processes of Ariadnaesporites sp., sample 1. (F) Palynomorph type A, sample 2, X 250. (G) Camarozonosporites sp., sample 1. (H) Palynomorph type B, sample 1. (I) Cavate spore (Balmeisporites-like), sample 1, X 450. (J) Cyathidites australis, sample 4. (K) Concavisporites sp., sample 2. (L) Fungal spore, sample 4. (M) Gabonisporis sp. cf. G. labyrinthus, sample 1. (N) Verrucosisporites sp., sample 4. (O) Ephedripites sp., sample 2 (SEM), X 700. (P) Steevesipollenites grambasti, sample 2 (SEM), X 1000. (Q) Balmeiopsis limbatus, sample 3. (R) Monosulcites sp., sample 3. (S and U) Pollen tetrads, sample 1 ((U) X 1200). (T) Dichastopollenites? sp. sensu Ibrahim and Abdel-Kireem, 1997; sample 1, X 1200. deposition. Also important is the state of preservation of palynofacies is dominated by large phytoclasts (mainly cuticular, and other particulate organic matter, and brown wood and some cuticles) and contain minor palynomorphs; the latter are part of the palynofacies amounts of poorly preserved phytoclasts and amorphous (Figs. 5–7). As has been stated in Section 4 the Bulaq organic matter (Fig. 4). Spores and pollen are diverse but 144 M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148

Fig. 6. (A) cf. Cretacaeiporites polygonalis, sample 2 (SEM), X 700. (B) Foveomonocolpites sp. 2, sample 2 (SEM), X 700. (C and O) Retimonocolpites sp. (Retimonocolpites Group sensu Herngreen, 1975) ((C) sample 2, SEM, X 700, (O) sample 1). (D) Oblique view of Proteacidites sp., sample 2, X 1200. (E) Palynomorph type C, sample 2 (SEM). (F) cf. Retidiporites magdalenensis, sample 2, X 700. (G) Syncolporites schrankii, sample 3. (H) cf. Stellatopollis sp., sample 2 (SEM), X 1000. (I) Arecipites sp., sample 3. (J and Q) Liliacidites spp. ((J) sample 4, X 1200, (Q) sample 2). (K) Foveotricolpites gigantoreticulatus, sample 3. (L) Monocolpopollenites spheroidites, sample 2. (M) Retidiporites? sp., sample 4. (N) Syncolporites subtilis, sample 4. (P) Foveomonocolpites sp. 1, sample 3.

show relatively small percentages compared to phyto- ynofacies is similar to an exinitic facies with spo- clasts and other organic debris. Fungal spores and fungal romorphs and abundant structured palynodebris hyphens are also present. Both the palynomorphs and (Habib, 1979). It is believed to have accumulated during the structured phytoclasts are well preserved. This pal- times of high sedimentation rates of land plant materials. M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 145

Fig. 7. (A) Longapertites sp., sample 3. (B) Foveomonocolpites sp. sensu Schrank, 1994a, sample 2. (C) Foveotricolpites tienabaensis, sample 2. (D and G) Foveotricolpites sp. cf. F. giganteus (D. sample 3, G. sample 2). (E) Retitricolporites fekelensis, sample 1. (F) Foveotricolpites gigantoreticulatus, sample 4. (H) Tricolpites microreticulatus, sample 4. (I) Proteacidites sp. sensu Schrank, 1994a, sample 1. (J) Proteacidites sp. 3 Lawal and Moullade, 1986; sample 2, X 700. (K) Foveotricolpites irregularis, sample 1. (L) Tricolpites maximus, sample 2. (M) Chomotriletes minor, sample 2. (N) Tricolpites sp., sample 2. (O) Momipites sp. sensu Schrank and Ibrahim, 1995; sample 2, X 1200. (P) Microforaminiferal test lining, sample 3. (Q) Tricolpites sp. S.CI. 278 Jardine and Magloire, 1965, sample 2, X 250. (R) Rhyzophagites sp., sample 2, X 300. (S) aﬀ. Celyphus sp., sample 2. (T) Pluricellaesporites sp., sample 2, X 450. (U) Ovoidites parvus, sample 4. (V) Ovoidites ligneolus, sample 4. (W) Botryococcus sp., sample 3.

Facies 1 of Firth (1993) is also similar and contains well- ported. Most land-derived organic matter, such as cuti- preserved indigenous phytoclasts and a small percentage cles, accumulates in fluvio-lacustrine sites close to source of miospores. Mukhopadhyay et al. (1985) found that vegetation (Batten, 1996). the source of well-preserved phytoclasts was a swampy Most cuticular remains of the Bulaq palynofa- environment. Freshwater conditions are therefore sup- cies debris are believed to have been derived from 146 M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 gymnosperms because their cuticles are thick (Batten, References 1975; Thomas, 1982). This is in spite of the fact that the palynofloras themselves are dominated by the angio- Aboul Ela, N.M., 1978. The palynological characteristics of the Upper sperms. This may also be attributed to the fact that Cretaceous sediments in Abu Tartur Plateau, Kharga Oasis, Egypt. Revista Espannola~ de Micropaleontologia 10, 421–442. angiosperm cuticles are less resistant and may be de- Adegoke, O.S., Jan Du Chêene, R.E., Agumanu, A.E., Ajayi, P.O., stroyed or winnowed away into more distant environ- 1978. Palynology and age of the Kerri-Kerri Formation, Nigeria. ments of deposition. Tracheids in this angiosperm- Revista Espannola~ De Micropaleontologia 10, 267–283. dominated association are rare, probably because an- Awad, G.H., Ghobrial, M.G., 1965. Zonal stratigraphy of the Kharga giosperms lack tracheids. However, most of the Permian Oasis. Egypt Geological Survey Paper 24, 77 p. Awad, M.Z., 1994. Stratigraphic, palynological and palaeoecological and Mesozoic wood is gymnospermous (Batten, 1996). studies in the east-central Sudan (Khartoum and Kosti Basins), Late Jurassic to Mid-Tertiary. Berliner geowissenschaftliche Abhandlungen A 161, 1–163. Azeema, C., Boltenhagen, E., 1974. Pollen de Creetace moyen du Gabon 6. Conclusions attribue aux Ephedrales. Paleebiologie Continentale 5, 1–37. Barthel, K.W., Herrmann-Degen, W., 1981. Late cretaceous and early tertiary stratigraphy in the Great Sand sea and its SE margins 1. The Bulaq palynofloras are exclusively derived from (Farafra and Dakhla Oases), SW Desert, Egypt. Mitteilungen terrestrial land plants. The occurence of abundant Bayerische Staatssammlung Palaaontologie€ Historische Geologie ferns (mainly Deltoidospora/Cyathidites) and the pres- 21, 141–182. ence of water ferns (Ariadnaesporites) and freshwater Batten, D.J., 1975. Wealden palaeoecology from the distribution of algae support the occurrence of widespread moist plant fossils. Proceedings of the GeologistsÕ Association, London 85, 433–458. conditions (water bodies and wet biotopes) under a Batten, D.J., 1996. Palynofacies. In: Jansonius, J., McGregor, D.C. warm-humid palaeoclimate. The palynomorphs and (Eds.), Palynology: Principles and Applications, American Asso- palynofacies suggest deposition in a fluvio-lacustrine ciation of Stratigraphic Palynologists Foundation 3, 1011–1064. palaeoenvironment close to source vegetation. Boltenhagen, E., 1967. Spores et pollen du Creetace supeerieur du 2. The previously recognized brackish palaeoenviron- Gabon. Pollen et Spores IX, 335–355. Boltenhagen, E., 1975. Pollen peripore du Creetace supeerieur du ment of the basal Quseir Formation in the Kharga Gabon. Revue de Micropaleeontologie 17 (4), 164–170. area could have been related to southward transgres- Boltenhagen, E., 1976. Pollen et spores Seenoniens du Gabon. Cahiers sion by an arm of the Tethys. de Micropaleeontologie, 150 p. 3. The palynological age of the basal part of the Quseir Boltenhagen, E., 1980. Palynologie du Creetace supeerieur du Gabon. Formation in the Bulaq area (Kharga Oasis) is Meemoirs de la Section des Sciences 7, 11–191. Collinson, M.E., 1991. Diversification of modern heterosporous Campanian (probably Early Campanian). This dat- pteridophytes. In: Blackmore, S., Barnes, S.H. (Eds.), Pollen and ing is based on certain marker angiosperm ranges Spores, Syst. Assoc. Spec. 44, Clarendon Press, Oxford, 119–150. such as Proteacidites sp. 3 Lawal and Moullade and Dettmann, M.E., 1963. Upper Mesozoic microfloras from south- Syncolporites schrankii, among others. eastern Australia. Proceedings of the Royal Society, Victoria 77 (1), 4. The Bulaq palynofloras are typical of the ‘‘Senonian 1–152. Dominik, W., 1985. Stratigraphie und Sedimentologie (Geochemie, Palmae Province’’ where Palmae and other protea- Schwermineralanalyse) der Oberkreide von Bahariya und ihre cean and syncolporate pollen occur. They are identi- Korrelation zum Dakhla-Becken (Western Desert, AAgypten).€ cal to their Senonian contemporary counterparts Berliner geowissenschaftlische Abhandlungen A 62, 173 p. from North Africa, but are significantly different El-Beialy, S.Y., 1994. Palynostratigraphy and palynofacies analysis of from those of West Africa and South America. some subsurface Cretaceous formations in the Badr El Dein (Bed 1- 1) borehole, North Western Desert, Egypt. Neues Jahrbuch fuur€ 5. Some Campanian-Maastrichtian pollen such as Echi- Geologie und Palaaontologie,€ Abhandlungen, 1994, 192 (2), pp. triporites, Spinizonocolpites and Periretisyncolpites 133–149. (the latter is an endemic African genus) are absent. El Beialy, S.Y., 1995. Campanian-Maastrichtian palynomorphs from This is probably because the basal Quseir association the Duwi (phosphate) formation of the Hamrawein and Umm El might be equivalent to an earlier Late Senonian paly- Hueitat mines, Red Sea, Egypt. Review of Palaeobotany and Palynology 85, 303–317. nofloral event. Firth, J.V., 1993. Palynofacies and thermal maturation analysis of sediments from the Nankai Trough. Proceedings of the Ocean Drilling Program, Scientific Results 131, p. 57–69. Acknowledgements Frederiksen, N.O., 1987. Tectonic and paleogeographic setting of a new Latest Cretaceous floristic province in North America. Palaios 5, 533–542. The author is indebted to the Egyptian General De- Germeraad, J.H., Hopping, C.A., Muller, J., 1968. Palynology of sert Development Organization (Management of the tertiary sediments from tropical areas. Review of Palaeobotany and ‘‘Groundwater’’ at Kharga) for making available the Palynology 6, 189–348. core material and the well logs. Thanks are also due to Habib, D., 1979. Sedimentology of palynomorphs and palynodebris in Cretaceous carbonaceous facies south of Vigo Seamount. Initial Mr. A. Abdel-Nasser (Assiut University) for operating Reports of the Deep Sea Drilling Project XLVII (2), pp. 451– the scanning electron microscope. 460. M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148 147

Hall, J.W., 1975. Ariadnaesporites and Glomerisporites in the Late Mahmoud, M.S., 1998. Palynological dating of the Quseir Formation, Cretaceous: ancestral Salviniaceae. American Journal of Botany Kharga Oasis (Egypt). Arab Gulf Journal of Scientific Research 62, 359–369. 16 (2), 267–281. Helal, A.H., 1965. Jurassic spores and pollen grains from the Kharga Mohsen, S.A., 1992. Cretaceous plant microfossils from the subsurface Oasis, Western Desert, Egypt. Neues Jahrbuch fuur€ Geologie und of Kharga Oasis, Western Desert, Egypt. Journal of African Earth Palaaontologie,€ Abhandlungen, 1965, 123 (2), 160–166. Sciences 14, 567–577. Hendriks, F., Luger, P., Kallenbach, H., Schroeder, J.H., 1984. Morgan, R., 1978. Albian to Senonian palynology of site 364, Angola Stratigraphical and sedimentological framework of the Kharga-Sin Basin. Initial Reports of the Deep Sea Drilling Project 40, 915– El Kaddab stretch (western and southern part of the Upper Nile 951. basin), Western Desert, Egypt. Berliner geowissenschaftliche Ab- Mukhopadhyay, P.K., Rullkootter,€ J., Schaeffer, R.G., Welte, D.H., handlungen A 50, 117–151. 1985. Facies and diagenesis of organic matter in Nankai Trough Hermina, M., 1990. The surroundings of Kharga, Dakhla and Farafra sediments, Deep Sea Drilling Project Leg 87A. Initial Reports oases. In: Said, R. (Ed.), The Geology of Egypt. Balkema, of the Deep Sea Drilling Project 87, 877–889. Rotterdam, pp. 259–292. Nichols, D.J., Ames, H.T., Traverse, A., 1973. On Arecipites Wode- Herngreen, G.F.W., 1975. Palynology of middle and upper Cretaceous house, Monocolpopollenites Thomson and Pflug, and the species strata in Brazil. Meded Rijks Geol. Dienst 26 (3), 39–91. ‘‘Monocolpopollenites tranquillus’’. Taxon 22, 241–256. Herngreen, G.F.W., Chlonova, A.F., 1981. Cretaceous microfloral Pacltova, B., 1961. On some plant microfossils from fresh-water provinces. Pollen et Spores 23, 441–555. sediments of the Upper Cretaceous (Senonian) in the south Herngreen, G.F.W., Kedves, M., Rovnina, L.V., Smirnova, S.B., 1996. Bohemian Basin. Sb. uustr. uust Geol. 26 (1959), 47–102. Cretaceous palynofloral provinces: a review. In: Jansonius, J., PetrosÕyants, M.A., Trofimov, D.M., 1971. Palynological characteris- McGregor, D.C. (Eds.), Palynology: Principles and Applications, tics of the Late Cretaceous of the Mali-Niger syneclise (Southern American Association of Stratigraphic Palynologists Foundation 3, Sahara). ByulletenÕ Moskovskogo Obshchestva Ispytatelei Prirody, 1157–1188. Otdelenie Geologija 76, 75–82 (in Russian). Ibrahim, M.I.A., Abdel-Kireem, M.R., 1997. Late Cretaceous palyno- Ravn, R.L., 1998. Taxon.exe, v 3.2 (Free Internet Database, http:// floras and foraminifera from Ain El-Wadi area, Farafra Oasis, www.bc.edu/bc_org/association/aasp/portal/ppaly.html). Egypt. Cretaceous Research 18, 633–660. Regali, M.S.P., Uesugui, N., Santos, A.S., 1974. Palinologia dos Ibrahim, M.I., Schrank, E., Abdel Kireem, M.R., 1995. Cretaceous sedimentos Meso-Cenozooicos do Brasil. Bol. Teec. Petrobraas 17 (4), biostratigraphy and palaeogeography of north Egypt and northeast 263–301. Libya. PRJ 7, 75–93. Said, R., 1962. The Geology of Egypt. Elsevier, Amsterdam. 377 p. Jan Du Chêene, R.E., 1977. Some new pollen species of the Salami, M.B., 1984. Cretaceous and Early Tertiary palynofacies of upper Maastrichtian, Tar Sand, Abeokuta Formation, south- south-western Nigeria. Revista Espannola~ de Micropaleontologia ern Nigeria. Revista Espannola~ de Micropaleontologia 9, 191–201. 16, 415–423. Jan Du Chêene, R.E., De Klasz, I., Archibong, E.E., 1978. Biostrati- Salami, M.B., 1985. Upper Senonian and Lower Tertiary pollen grains graphic study of the borehole Ojo-1, SW Nigeria, with special from the Southern Nigeria Sedimentary Basin. Revista Espannola~ de emphasis on the Cretaceous microflora. Revue de Micropaleeon- Micropaleontologia 17, 5–26. tologie 2 (3), 123–139. Salami, M.B., 1990. Palynomorph taxa from the Lower Coal Jardinee, S., Magloire, L., 1965. Palynologie et stratigraphie du Creetace Measures deposits (?Campanian-Maastrichtian) of Anambra des Bassins du Seeneegal et de Coote^ dÕIvoire. Meemoires Bureau Trough, southeastern Nigeria. Journal of African Earth Sciences Recherches Geeologiques Minieeres 32, 187–245. 11, 135–150. Kaska, H.V., 1989. A spore and pollen zonation of Early Cretaceous Salard-Cheboldaeff, M., 1990. Intertropical African palynostratigra- to Tertiary nonmarine sediments of Central Sudan. Palynology 13, phy from Cretaceous to Late Quaternary times. Journal of African 79–90. Earth Sciences 11 (1/2), 1–24. Klitzsch, E., Hermina, M., 1989. The mesozoic. In: Hermina, M., Schrank, E., 1984. Organic-geochemical and palynological studies of a Klitzsch, E., List, K. (Eds.), Stratigraphic Lexicon and Explana- Dakhla Shale profile (Late Cretaceous) in southeast Egypt. Berliner tory Notes to the Geological Map of Egypt 1:500 000. Conoco Inc. Geowissenschaftliche Abhandlungen A 50, 189–207. and E.G.P.C., Cairo, pp. 77–139. Schrank, E., 1987a. Biostratigraphic importance of microfloras from Klitzsch, E., Squyres, C.H., 1990. Paleozoic and Mesozoic geological the Late Cretaceous clastic series of northwestern Sudan. Creta- history of northeastern Africa based upon new interpretation of ceous Research 8, 29–42. Nubian strata. American Association of Petroleum Geologists Schrank, E., 1987b. Palaeozoic and Mesozoic palynomorphs from Bulletin 74, 1203–1211. northeast Africa (Egypt and Sudan) with special reference to Late Klitzsch, E., Harms, J.C., Lejal-Nicol, A., List, F.K., 1979. Major Cretaceous pollen and dinoflagellates. Berliner Geowissenschaftli- subdivisions and depositional environments of Nubia Strata, che Abhandlungen A 75.1, 249–310. southwestern Egypt. American American Association of Petroleum Schrank, E., 1994a. Palynology of the Yesomma Formation in the Geologists Bulletin 63, 967–974. northern Somalia: a study of pollen, spores and associated Klitzsch, E., List, F.K., Poohlmann,€ G. (Eds.), 1987. Geological map of phytoplankton from the Late Cretaceous Palmae Province. Pala- Egypt 1:500,000, Sheet NG 35 SE Dakhla. Conoco Coral, Egyptian eontographica B 231, 63–112. General Petroleum Corporation, Cairo. Schrank, E., 1994b. Nonmarine Cretaceous palynology of northern Kuyl, O.S., Muller, J., Waterbolk, H.T., 1955. The application of Kordofan, Sudan, with notes on fossil Salviniales (water ferns). palynology to oil geology, with special reference to western Geologische Rundschau 83, 773–786. Venezuela. Geologie en Mijnbouw 17 (3), 49–75. Schrank, E., Ibrahim, M.I.A., 1995. Cretaceous (Aptian-Maastrich- Lawal, O., 1982. Biostratigraphie, palynologique et paleeo-environne- tian) palynology of foraminifera-dated wells (KRM-1, AG-18) in ments des formations Creetaceees de la Haute-Benoue, Nigeria, nord- northwestern Egypt. Berliner Geowissenschaftliche Abhandlungen oriental. Theese de Doctorat de Speecialitee(3c cycle), Universiteede A 177, 1–44. Nice, 198 p. Schrank, E., Mahmoud, M.S., 1998. Palynology (pollen, spores Lawal, O., Moullade, M., 1986. Palynological biostratigraphy of and dinoflagellates) and Cretaceous stratigraphy of the Dakhla Cretaceous sediments in the upper Benue Basin, N.E. Nigeria (1). Oasis, central Egypt. Journal of African Earth Sciences 26 (2), 167– Revue de Micropaleeontologie 29, 61–83. 193. 148 M.S. Mahmoud / Journal of African Earth Sciences 36 (2003) 135–148

Schrank, E., Mahmoud, M.S., 2000. New taxa of angiosperm pollen, Srivastava, S.K., 1977. Miospores from the Frederecksburg Group miospores and associated palynomorphs from the early Late (Albian) of the southern United States. Paleebiologie continentale Cretaceous of Egypt (Maghrabi Formation, Kharga Oasis). (1975) 6 (2), 1–119. Review of Palaeobotany and Palynology 112, 167–188. Sultan, I.Z., 1985. Palynological studies in the Nubia Sandstone Schrank, E., Mahmoud, M.S., 2002. Barremian angiosperm pollen and Formation, east of Aswan, southern Egypt. Neues Jahrbuch fuur€ associated palynomorphs from the Dakhla Oasis area, Egypt. Geologie und Palaaontologie,€ Monatshefte 10, 605–617. Palaeontology 45 (1), 33–56. Tea-Yassi, J., Digbehi, Z.B., Yao, K.R., Glohi, B.V., 1999. EEtude de Schrank, E., Nesterova, E.V., 1993. Palynofloristic changes and quelques palynomorphes du Creetace Supeerieur du bassin de Coote^ Cretaceous climates in northern Gondwana (NE Africa) and dÕIvoire: implications biostratigraphiques et paleeoenvironn- southern Laurasia (Kazakhstan). In: Thorweihe, U., Schandel- mentales. Journal of African Earth Sciences 29(4), 783–798. meier, H. (Eds.), Geoscientific Research in Northeast Africa. Thomas, B.M., 1982. Land-plant source rocks for oil and their Balkema, Rotterdam, pp. 381–390. significance in Australian basins. The APEA Journal (Australian Singh, C., 1971. Lower Cretaceous microfloras of the Peace River area, Petroleum Exploration Association) 22, 164–178. northwestern Alberta. Research Council of Alberta Bulletin 28 (2), Thomson, P.W., Pflug, H., 1953. Pollen und Spores des Mitteleuro- 542. paischen Tertiaars.€ Palaeontographica B 94, 1–138. Smith, A.G., Hurley, A.M., Briden, J.C., 1982. Palaaokontinentale€ Van Erve, A., Mohr, B., 1988. Palynological investigations of the Late Weltkarten des Phanerozoikums. Stuttgart (Enke), 102 p. Jurassic microflora from the vertebrate locality Guimarota coal Soliman, H.A., 1977. Foraminifeeres et microfossiles veegeetaux prove- mine (Leiria, central Portugal). Neues Jahrbuch fuur€ Geologie und nant du ‘‘Nubia Sandstone’’ de subsurface de LÕOasis El Kharga, Palaaontologie,€ Monatshefte 4, 246–262. Deesert de LÕOuest, Egypte. Revue de Micropaleeontologie 20, 114– Youssef, M.I., 1957. Upper Cretaceous rocks in Kosseir area. Inst. 124. Bulletin of the Egyptian Desert Institute 7, 35–53.