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Middle Eocene dinoflagellate cysts from the Tsoul section, eastern External Rif, Morocco: biostratigraphy and paleoenvironmental interpretations

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This palynological study is carried out on the Eocene sediments in the Tsoul section, which is located in the Taza region, Tsoul Unit, eastern External Rif, northeastern Morocco. It aims their age assessment and paleoenvironmental interpretations, based on palynological data. The palynological content is diverse, well preserved and rich in organic-walled dinoflagellate cysts (dinocysts), whereas spores and pollen are rare. We identified 117 dinocyst species (including some reworked species), among which 6 species and 1 acme are good biostratigraphic markers of the Bartonian (middle Eocene). The main dinocyst bioevents that are used to assign the studied section to the Bartonian include the lowest occurrences (LOs) of Glaphyrocysta semitecta, Impagidinium dispertitum, Lentinia serrata, Membranophoridium aspinatum and Operculodinium divergens, the highest occurrence (HO) of Castellodinium compactum and the acme of Polysphaeridium spp. Regarding the paleoenvironment, the studied section was deposited in a marine environment, characterized by a low productivity inferred from the scarcity of peridinioid cysts and by important environmental changes ranging from restricted inner neritic (lagoon) conditions, inferred from the dominance of Homotryblium and Polysphaeridium, to outer neritic open marine conditions inferred from the dominance of Spiniferites.

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The authors thank the Department of Earth Science and the Laboratory of Geo-Biodiversity and Natural Patrimony (“Geophysics, Natural Patrimony and Green Chemistry” Research Center), Scientific Institute, for the technical help. A. Toufiq (University Chouaïb Doukkali, El Jadida, Morocco) is also thanked for his help during the field work. This study is a part of the first author (I. Mahboub)’s PhD thesis, directed by H. Slimani. The journal Editor-in-Chief Abdullah M. Al-Amri, the Chief Editor Beatriz Bádenas, the Associate Editor Idoia Rosales, and the reviewers Przemysław Gedl and two anonymous are thanked for their corrections, comments and suggestions that improved considerably the manuscript. We are very grateful for their help.

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Correspondence to Hamid Slimani.

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Responsible Editor: Idoia Rosales



List of dinoflagellate cyst taxa and other palynomorphs (spores, pollen, foraminifera linings) observed in the Tsoul section (eastern External Rif, northeastern Morocco). The dinocyst taxa are arranged alphabetically by genera order in Appendix, and by their first occurrence order in Table 1. The nomenclature of the dinocyst species and references to their authors follow DINOFLAJ3 (Williams et al. 2017)

Achilleodinium biformoides (Eisenack 1954) Eaton, 1976.

Achomosphaera alcicornu (Eisenack 1954) Davey and Williams 1966.

Achomosphaera minor He Chengquan 1991.

Adnatosphaeridium vittatum Williams and Downie 1966.

Alterbidinium bicellulum (Islam 1983) Lentin and Williams 1985.

Alterbidinium distinctum (Wilson 1976) Lentin & Williams 1976.

Alterbidinium earnleyense (Islam 1983) Lentin and Williams 1985. (Plate 6: 1).

Amphorosphaeridium multispinosum (Davey and Williams 1966) Sarjeant 1981.

Apectodinium homomorphum (Deflandre and Cookson 1955) Lentin and Williams 1977.

Apteodinium australiense (Deflandre and Cookson 1955) Williams 1978. (Plate 1: 18).

Apteodinium reticulatum Singh 1971.

Areoligera medusettiformis Wetzel 1933.

Areoligera senonensis Lejeune-Carpentier 1938.

Ataxiodinium choane Reid, 1974.

Batiacasphaera compta Drugg, 1970b. (Plate 2: 3, 4).

Bitectatodinium sp. A of Mahboub et al., 2019.

Castellodinium compactum (Michoux 1988) Williams et al. 2015. (Plate 3: 7).

Cerodinium speciosum (Alberti 1959) Lentin and Williams 1987.

Charlesdowniea coleothrypta (Williams and Downie 1966) Lentin and Vozzhennikova 1989. (Plate 3: 3, 6).

Chiropteridium cf. conispinum Williams 1978.

Chiropteridium galea (Maier 1959) Sarjeant 1983.

Circulodinium distinctum (Deflandre and Cookson 1955) Jansonius 1986.

Cleistosphaeridium ancyreum (Cookson and Eisenack 1965) Eaton et al. 2001.

Cleistosphaeridium diversispinosum Davey et al. 1966.

Cleistosphaeridium polypetellum (Islam 1983) Stover and Williams 1995.

Cordosphaeridium delimurum Fensome et al. 2009.

Cordosphaeridium fibrospinosum Davey and Williams 1966.

Cribroperidinium boreas (Davey, 1974) Helenes 1984.

Cribroperidinium edwardsii (Cookson and Eisenack 1958) Davey 1969.

Cribroperidinium giuseppei (Morgenroth 1966) Helenes 1984.

Cribroperidinium tenuitabulatum (Gerlach 1961) Helenes 1984.

Danea impages Damassa 1979. (Plate 2: 19, 20).

Deflandrea granulata Menéndez 1965.

Deflandrea phosphoritica Eisenack 1938.

Diphyes bifidum Antolinez-Delgado and Oboh-Ikuenobe 2007. (Plate 2: 7, 8).

Diphyes colligerum (Deflandre and Cookson 1955) Cookson 1965.

Duosphaeridium nudum (Cookson 1965) Loeblich Jr. and Loeblich III 1968. (Plate 2: 5, 6).

Emmetrocysta urnaformis (Cookson 1953) Stover 1975.

Enneadocysta arcuata (Eaton 1971) Stover and Williams 1995. (Plate 2: 16, 17).

Fibrocysta vectensis (Eaton 1976) Stover and Evitt 1978. (Plate 2: 21).

Florentinia radiculata (Davey and Williams 1966) Davey and Verdier 1973.

Gelatia inflata Bujak 1984.

Glaphyrocysta exuberans (Deflandre and Cookson 1955) Stover and Evitt 1978.

Glaphyrocysta inculta (Morgenroth 1966b) Stover and Evitt 1978.

Glaphyrocysta intricata (Eaton 1971) Stover and Evitt 1978.

Glaphyrocysta retiintexta (Cookson 1965) Stover and Evitt 1978.

Glaphyrocysta semitecta (Bujak in Bujak et al. 1980) Lentin and Williams 1981. (Plate 3: 8, 9).

Glaphyrocysta vicina (Eaton 1976) Stover and Evitt 1978.

Aff. Heslertonia heslertonensis of Heilmann-Clausen and Costa, 1989 (Plate 1: 15, 16)

Homotryblium abbreviatum Eaton 1976. (Plate 2: 14, 15).

Homotryblium tenuispinosum Davey and Williams 1966.

Hystrichokolpoma bullatum Wilson 1988. (Plate 2: 10, 11).

Hystrichokolpoma globulus Michoux 1985.

Hystrichokolpoma rigaudiae Deflandre and Cookson 1955. (Plate 2: 12).

Hystrichokolpoma salacia Eaton 1976. (Plate 2: 9).

Hystrichokolpoma truncatum Biffi and Manum 1988.

Hystrichokolpoma unispinum Williams and Downie 1966.

Hystrichosphaeropsis ovum Deflandre 1935. (Plate 1: 12, 13).

Ifecysta lappacea (Drugg 1970) Antolinez-Delgado and Oboh-Ikuenobe 2007. (Plate 1: 17; Plate 2: 22).

Impagidinium aspinatum (Cookson and Eisenack 1974) Damassa 1979. (Plate 1: 8).

Impagidinium dispertitum (Cookson and Eisenack 1965) Stover and Evitt 1978. (Plate 1: 10, 11).

Impagidinium maculatum (Cookson and Eisenack 1961b) Stover and Evitt 1978.

Impagidinium margaritiferum (Cookson and Eisenack 1960) Stover and Evitt 1978.

Impagidinium simplicium (Cookson and Eisenack 1961) Stover and Evitt 1978.

Isabelidinium pellucidum (Deflandre and Cookson 1955) Lentin and Williams 1977.

Kallosphaeridium yorubaense Jan du Chêne and Adediran 1985. (Plate 2: 18).

Kenleyia lophophora Cookson and Eisenack1965.

Lejeunecysta beninensis Biffi and Grignani 1983. (Plate 1: 5).

Lejeunecysta hyalina (Gerlach 1961) Artzner and Dörhöfer 1978.

Lentinia serrata Bujak et al. 1980. (Plate 1: 14).

Lentinia wetzelli (Morgenroth 1966) Bujak in Bujak et al. 1980. (Plate 1: 7).

Lingulodinium machaerophorum (Deflandre and Cookson 1955) Wall 1967.

Manumiella seelandica (Lange 1969) Bujak and Davies 1983.

Melitasphaeridium pseudorecurvatum (Morgenroth 1966) Bujak et al. 1980.

Membranophoridium aspinatum Gerlach 1961. (Plate 3: 1, 2).

Mendicodinium brunneum Bucefalo Palliani et al., 1997.

Microdinium setosum Sarjeant, 1966.

Muratodinium fimbriatum (Cookson and Eisenack 1967) Drugg 1970.

Nelsoniella tuberculata Cookson and Eisenack 1960.

Odontochitina costata Alberti 1961.

Oligosphaeridium complex (White 1842) Davey and Williams 1966.

Oligosphaeridium granulatum He Chengquan, 1991.

Operculodinium centrocarpum (Deflandre and Cookson 1955) Wall 1967.

Operculodinium divergens (Eisenack 1954) Stover and Evitt 1978.

Operculodinium nanaconulum Islam 1983. (Plate 2: 1, 2).

Operculodinium placitum Drugg and Loeblich Jr. 1967.

Palaeocystodinium golzowense Alberti 1961.

Palaeohystrichophora palaeoinfusa Fensome et al. 2009.

Pentadinium laticinctum Gerlach 1961.

Phelodinium magnificum (Stanley 1965) Stover and Evitt 1978.

Phthanoperidinium alectrolophum Eaton 1976. (Plate 1: 4).

Phthanoperidinium cf. levimurum Bujak et al. 1980.

Phthanoperidinium stockmansii (De Coninck 1975) Lentin and Williams 1977 (Plate 1: 1).

Polysphaeridium biformum Islam 1983. (Plate 2: 13).

Polysphaeridium subtile Davey and Williams 1966.

Pterodinium cingulatum (Wetzel 1933) Below 1981. (Plate 1: 9).

Pterodinium sp. cf. Pentadidium polypodum Edwards 1982.

Pyxidinopsis elliptica Biffi and Manum 1988.

Riculacysta perforata Stover 1977.

Samlandia chlamydophora Eisenack 1954. (Plate 1: 19).

Schematophora obscura Wilson 1988.

Selenopemphix nephroides Benedek 1972. (Plate 1: 2).

Senegalinium cf. orei (Jan du Chêne and Adediran 1985) Stover and Williams 1987.

Senegalinium laevigatum (Malloy 1972) Bujak and Davies 1983.

Sentusidinium sparsibarbatum Erkmen and Sarjeant 1980.

Spinidinium densispinatum Stanley 1965.

Spinidinium essoi Cookson and Eisenack 1967. (Plate 1: 3).

Spiniferella cornuta (Davey and Williams 1966) Williams et al. 1998.

Spiniferites membranaceus (Rossignol 1964) Sarjeant 1970.

Spiniferites ramosus (Ehrenberg 1837) Mantell 1854.

Spiniferites twistringiensis (Maier 1959) Fensome et al. 1990.

Stenodinium meckelfeldense (Gocht 1969) Williams et al. 2015 Stover and Williams 1995.

Tectatodinium pellitum Wall 1967.

Thalassiphora pelagica (Eisenack 1954) Eisenack and Gocht 1960.

Vozzhennikovia apertura (Wilson 1967) Lentin and Williams 1976.

Wetzeliella articulata Eisenack 1938. (Plate 3: 4, 5).

Ynezidinium brevisulcatum (Michoux 1985) Lucas-Clark and Helenes 2000.

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Mahboub, I., Slimani, H. Middle Eocene dinoflagellate cysts from the Tsoul section, eastern External Rif, Morocco: biostratigraphy and paleoenvironmental interpretations. Arab J Geosci 13, 197 (2020). https://doi.org/10.1007/s12517-020-5165-7

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  • Middle Eocene
  • Dinocysts
  • Biostratigraphy
  • Paleoenvironment
  • Tsoul section
  • Northeastern Morocco