Cretaceous Research 106 (2020) 104219

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Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

Dinoflagellate cyst evidence for the age, palaeoenvironment and

paleoclimate of a new CretaceousePaleogene (K/Pg) boundary section
at the Bou Angueur syncline, Middle Atlas, Morocco
Sara Chakir a, b, Hamid Slimani b, *, Touria Hssaida a, La

szlo

Kocsis c,
Emmanuel Gheerbrant , Nathalie Bardet , Nour-Eddine Jalil d, Mustapha Mouflih a,
d d

Imane Mahboub b, Hassan Jbari b

a
Laboratory of Dynamics of Sedimentary Basins and Geological Correlations, Department of Geology, Faculty of Sciences Ben M'Sik, Hassan II University of
Casablanca, Avenue Driss El Harti, BP 7955, Sidi Othmane, 20800, Casablanca, Morocco
b
Geo-Biodiversity and Natural Patrimony Laboratory (GEOBIO), “Geophysics, Natural Patrimony and Green Chemistry” Research Center (GEOPAC),
Scientific Institute, Mohammed V University in Rabat, Avenue Ibn Batouta, P.B. 703, 10106, Rabat-Agdal, Morocco
c
Faculty of Science, Jalan Tungku Link, Gadong, Universiti Brunei Darussalam, Brunei Darussalam, BE141, Indonesia
d ^timent de pal

Entre sol aile Valhulbert ba eontologie, 8 rue Buffon, 75005, Paris, France

a r t i c l e i n f o a b s t r a c t

Article history: In this paper, we present results of a palynological analysis of the phosphate succession from the
Received 26 February 2019 northern flank of the Bou Angueur syncline in the Middle Atlas (central Morocco). The studied interval
Received in revised form yielded a well-preserved and diverse palynological content, dominated by dinoflagellate cysts (dino-
16 July 2019
cysts), allowing a detailed taxonomic study of dinocyst and a new biostratigraphic dating of the section.
Accepted in revised form 13 August 2019
Available online 24 August 2019
The studied section is assigned to the upper MaastrichtianeDanian interval. A new, relatively complete
K/Pg boundary transition was recognized. The succession at the Bou Angueur syncline was compared to
Tethyan K/Pg boundary sections, such as the K/Pg boundary Global Stratotype Section and Point (GSSP) at
Keywords:
Dinocysts
El Kef (NW Tunisia) and Ouled Hadou section (NE Morocco). The dinocyst marker events used for the
K/Pg boundary identification of the late Maastrichtian include the first occurrences of Cordosphaeridium inodes subsp.
Palynostratigraphy inodes, Deflandrea galeata and Glaphyrocysta perforata and those used for the early Danian include the
Paleoenvironment first occurrences of Carpatella cornuta, Cassidium fragile, Danea californica, Lanternosphaeridium rein-
Paleoclimate hardtii and Senoniasphaera inornata.
NW Africa The organic-walled dinocyst-producing dinoflagellates did not undergo the K/Pg mass extinction
resulting from a large meteoritic impact. They enable us to interpret the environmental and climatic
changes in the earliest Danian, while calcareous microfossils could not, since they are partially or
completely extinct. We thus describe seven palynological assemblages, reflecting paleoenvironmental
and paleoclimatic changes, inferred from changes in relevant ecological parameters, including changes in
the relative abundance of selected dinocyst groups, often used as paleoenvironmental indicators. An
inner to middle neritic marine environments alternated with several short phases of more offshore
conditions is deduced for the late Maastrichtian and earliest Danian and an open outer neritic marine
environment during a transgressive regime in the latest Danian. Several dinocyst events were recognized
in the studied interval, among which remarkable blooms of tropical warm-water and high-latitude cold-
water dinocyst taxa are interpreted herein as ecological responses, related to global climatic changes,
including a global warming episode during the late Maastrichtian and the well-known brief global
cooling following the K/Pg boundary Chicxulub impact.
© 2019 Elsevier Ltd. All rights reserved.

1. Introduction

The CretaceousePaleogene (K/Pg) boundary (~66 Ma) mass

* Corresponding author. Fax: þ212 537774540.
E-mail addresses: slimani@israbat.ac.ma, h_slimani@yahoo.com (H. Slimani).
extinction is the best-known crisis of Earth history among the

https://doi.org/10.1016/j.cretres.2019.104219
0195-6671/© 2019 Elsevier Ltd. All rights reserved.
2 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
public, as it concerns the extinction of the non-avian dinosaurs and
about 50e75% of all terrestrial and marine taxa. In the oceans, this
crisis has influenced the food chain and caused almost a complete
extinction (99%) of planktonic foraminifera (Keller, 1988, 2001;
Zachos et al., 1989; Kump, 1991, 2003; MacLeod et al., 1997;
Molina et al., 1998; Keller et al., 2002; Luciani, 2002). The causes
of this mass extinction are still subject to debate. Some researchers
advocate the catastrophic impact of a large meteorite in Chicxulub
(Yucatan peninsula, Mexico) (Alvarez et al., 1980; Hildebrand et al.,
1991; Gradstein et al., 2004; Molina et al., 2006; Schulte et al.,
2010), however others evoke the influence of intense volcanism
(eg. Deccan volcanism in India) that could be considered as the
catalyst, leading to environmental changes responsible of the K/Pg
boundary mass extinction (McLean, 1985; Courtillot et al., 1986,
1988; Courtillot, 1990; Punekar et al., 2014; Wilson, 2014; Wilson
et al., 2014; Adatte, 2015; Font et al., 2016, 2018; Keller et al.,
2016, 2018).
Previous paleoenvironmental and paleoclimatic studies of the
K/Pg boundary Chicxulub impact suggest a brief period of global
cooling called “impact winter”, caused by a thick cloud of sulphate
aerosols and soot as well obscuring the sky and blocking the sun-
light (Alvarez et al., 1980; Pierazzo et al., 2003; Vellekoop et al.,
2014, 2015, 2016). This period is followed by a period of green-
house warming caused by CO2 released into the atmosphere by the
impact (Kring, 2007; Vellekoop et al., 2018). A recent study
combining organic-walled dinoflagellate cyst and benthic fora-
minifer analyses across the K/Pg boundary of a number of Tethyan
sections, enabled to recognize four phases: the Maastrichtian or
pre-impact phase, a disaster phase at the K/Pg boundary, followed
by a recovery phase and an early Paleocene phase (Vellekoop et al.,
2017).
The K/Pg Boundary Stratotype and Global Stratotype Section and
Point has been defined at El Kef (Tunisia) (International Union of
Geological Sciences (IUGS), 1991; Molina et al., 2006) based on the
foraminifer mass extinction and on the thin clay layer with an
anomalous high iridium concentration that characterize this
boundary (Alvarez et al., 1980; Schulte et al., 2010). The dinofla-
gellate cysts e the main tool of our study e have allowed the age
determination of the K/Pg boundary at El Kef, and in contrast to
calcareous foraminifera, they did not undergo a mass extinction
(Brinkhuis and Zachariasse, 1988; M'Hamdi et al., 2015). Similar
evidences were recorded at many K/Pg boundary extinctions
(Hansen, 1977; De Coninck and Smit, 1982; Firth, 1987; Habib et al.,
1996; M'Hamdi et al., 2013; Mohamed et al., 2012; Açıkalın et al.,
2015; Slimani et al., 2010, 2016; Taba ra
 and Slimani, 2016;
Vellekoop et al., 2015, 2016; Taba ra
 et al., 2017; Soliman and
Slimani, 2019).
In Morocco, palynological studies, dealing with the systematics
of dinoflagellate cysts, biostratigraphy and paleoenvironment
across the K/Pg boundary, are rare. Most of them were carried out in
the Atlantic deposits of the “phosphate Plateau” (Doubinger, 1979;
Pre
vo^ t et al., 1979; Rauscher and Doubinger, 1982; Rauscher, 1985;
Soncini and Rauscher, 1988; Soncini, 1990) and in the Tethyan de-
posits of the external Rif Chain (Slimani et al., 2008, 2010, 2012,
2016; Slimani and Toufiq, 2013; Gue
de

et al., 2014; Gue

de

, 2016).
In the Middle Atlas, biostratigraphic and paleoenvirnmental studies
of the CretaceousePaleogene interval in the Bou Angueur syncline,
based on dinoflagellate cysts, spores and pollen (Herbig and
Fechner, 1994) and in the Oudiksou syncline based on char-
ophytes (Mebrouk et al., 2009), are preliminary and do not provide
detailed information about the completeness of the K/Pg boundary
interval. According to the latter authors, the study and the dating of
this interval have been proven to be very complex because of the
numerous lateral facies changes caused by pre- and syn-
sedimentary vertical movements.
Our study, carried out in the northern flank of the Bou Angueur
syncline in the Middle Atlas Mountains (Morocco) (Fig. 1), is a
continuation of our previous palynological studies undertaken as
part of our research project on the CretaceousePaleogene bound-
ary in Morocco. It aims to establish a high resolution dinocyst
stratigraphy of the CretaceousePaleocene transition, to refine the
age of this succession and identify the K/Pg boundary, to provide
paleoenvironmental and paleoclimatic reconstructions and to
compare the results with those of other K/Pg boundary sections,
including the GSSP at El Kef (Tunisia) and the Ouled Haddou section
(Morocco).
2. Geological settings
The Middle Atlas, where the study section is located, is an intra-
continental chain, which belongs to the Atlas system and repre-
sents a main vestige of the alpine orogeny in Morocco. It extends
from the southwest to the northeast over a distance of 350 km. It is
bordered to the west and the east by the Mesetian Domains, to the
south by the Central High Atlas Chain and to the north by the Rif
Chain. It is a result of the convergence between Europe (Iberia) and
Africa starting in the Late Cretaceous, and the culmination during
the MioceneePliocene times (Brede et al., 1992). Previous paleo-
ecological studies in the Middle Atlas suggest that the Jurassic
marine deposits belong to the Tethyan Ocean, while the Upper
Cretaceousemiddle Eocene deposits belong to the Atlantic Ocean,
with a long period of emersion during the Late Jurassic and the
beginning of the Early Cretaceous (see Frizon de Lamotte et al.,
2008). The general east-west transgression of the Atlantic Ocean
towards the Tethyan domain begun in the Turonian (Charrie re,
1991, 1992) and from the Santonian, the Middle Atlas was con-
nected to the Atlantic through the “phosphate Plateau” (Frizon de
Lamotte et al., 2008).
The Middle Atlas is subdivided into two different structural
units (Piquet, 2007): the folded Middle Atlas to the southeast and
the tabular Middle Atlas to the northwest. The two units are
separated by a major northeast-southwest trending fault eThe
main structural feature in the Middle Atlas Chaine called the
Northern Middle Atlas Fault (NMAF). The NMAF has played a very
important role in the Paleogene and Neogene sedimentation, rep-
resenting a boundary between a domain that includes a Cretaceous
series and a domain without it (Charrie re, 1990).
The folded Middle Atlas, where the study section is located,
corresponds to the highest reliefs (Bounacer mount: 3340 m high
and O-Salah mount: 3190 m high). It consists essentially of four
anticlines whose north and west boundaries are limited by major
faults that are locally transformed into ridges. These anticlines are
separated by large synclines, whose geometry and lithology are
controlled essentially by the remobilization of NMAF.
In the southern part of the folded Middle Atlas, the Upper
Cretaceous and Tertiary deposits are found in basins or synclines in
the central part of the mountain chain (Herbig, 1991), whose the
most known are:
1) the Bekrit syncline (also called El Koubbat syncline) belongs to
the southernmost part of the Middle-Atlas and is located in the
north of the NMAF;
2) the Bou Angueur syncline e our study area e is separated from
the previous syncline by the NMAF and the connected anticline
(Colo, 1961);
3) the Feleddi syncline (also called Ain Nokra syncline) is located in
the northeast of the Bou Angueur syncline;
4) the Oudiksou syncline is separated from the Feleddi syncline by
the northeast-southwest Middle Atlas Fault and its accompa-
nying anticline (Colo, 1961).
 S. Chakir et al. / Cretaceous Research 106 (2020) 104219 3

Fig. 1. Location and geological setting of the CretaceousePaleogene boundary section of Bou Angueur, Middle Atlas, Morocco. (B) is adapted from the geological map of Morocco
1:1000,000.

3. Stratigraphic overview of the Upper Cretaceous and lower
Paleocene in the Bou Angueur syncline
The Bou Angueur syncline is a northeast-southwest asymmet-
rical structure that belongs to the folded Middle Atlas. It contains a
post-Jurassic succession, which starts from the Cenomanian at its
southern flank and from the upper Campanian to Maastrichtian at
its northern flank (Termier, 1936; Herbig, 1988). It is further
distinguished by the most complete Paleogene and Neogene suc-
cession preserved in the Middle Atlas (west of Boulemane) and by
extensive Quaternary lava flows, which cover mostly its central and
southwestern parts (Dresnay, 1969; Martin, 1973, 1981; Herbig,
1993).
In the southern flank of the Bou Angueur syncline, the Creta-
ceous deposits start with the Lower Cretaceous succession of the

rie rose
“se
” and “se

rie jaune” of the Sidi Larbi Formation (Andreu
4 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
and Charriere, 1986). This succession is followed by the calcareous
sediments of the Amghourzif Formation (Andreu and Charrie re,
1986), which were assigned to the upper Cenomanian to lower
Turonian by Lehman and Herbig (2009), based on ammonites. The
Amghourzif Formation is overlain by the succession of Coniancian
to Santonian marls and siltstone of the Ait Sba’ Formation (Andreu
and Charriere, 1986) and Thanetian to upper Lutetian limestones of
the Bekrit-Timahdith Formation (Herbig, 1991, 1993), with a gap of
Campanian to Danian deposits of the Ait Sba’ Formation.
In the northern flank of the Bou Angueur syncline, where our
study section is located, the upper part of the Ait Sba’ Formation is
strongly diachronous and grades into the Campanian?eMaas-
trichtian to lower Thanetian of the Foum Kheneg Formation. The
Foum Kheneg Formation rests with an angular unconformity into
Middle Liassic (Domerian) limestones. This formation contains all
Campanian and Maastrichtian bituminous and phosphate deposits
(limestones, marls, bioclastic phosphorites, sandstones) (Ensslin;
1993) and the equivalent of Danian to lowermost Thanetian red
beds of the Irbzer Formation (Herbig and Fechener, 1994). The
Irbzer Formation is widespread elsewhere in other synclines in the
Middle Atlas, such as the synclines of Bekrit, Oudiskou and Felddi
(Rahhali, 1970; Charriere, 1990; Herbig, 1991; Herbig and Fechner,
1994). The Foum Kheneg Formation is surmounted by 28 m thick
of white and light rose-colored, thick-bedded bioclastic limestones,
similar to those encountered in the Bekrit-Timahdith synclines
(Herbig, 1991; Herbig and Fechner, 1994). Herbig and Fechner
(1994) undertook a palynological study, suggesting the upper
Danianelower Thanetian interval for the Foum Kheneg Formation
in the Bou Anguer syncline, based on the dominance of the dinocyst
Areoligera spp.
4. Paleoenvironmental and paleogeographic setting
During the Late Cretaceous and Palaleogne, the Middle Atlas
area constitutes a portion of a platform connecting the Atlantic and
Tethyan domains (Choubert, 1948; Michard, 1976). Previous studies
in this area have only highlighted the presence of phosphate and
bituminous deposits of the CampanianeEocene succession, and
little information is available on the environmental conditions of
these deposits, including the phosphate deposits of the Foum
Kheneg Formation, analyzed in our study. Phosphate and bitumi-
nous sediments have been deposited in tilted block basins that
emphasize the importance of the Northern Middle Atlas and
Southern Middle Atlas Faults and their adjacent structures (Mouflih
et al., 2006).
According to Mouflih et al. (2006), the Upper Cretaceouse
Paleogene succession is deposited in an epicontinental sea less
than 500 m deep. The phosphate deposits contain a high amount of
silts and marls and an important Maastrichtian macrofauna of fish,
reptiles and mollusks, which are reworked from the high infratidal
environment. The P2O5 content is significant, but does not exceed
26% and the sedimentation was slow with an important continental
influence, according to petrographic and mineralogical analyzes.
However, the bituminous deposits are rich in marine amorphous
organic matter and dinocysts. Taking into account previous studies
(Slansky, 1980; Belayouni, 1983; Aït Slimane, 1989; Herbig, 1991),
Mouflih et al. (2006) consider that the organic matter-phosphate
relationship is conditioned by an oxidizing redox potential with
respect to the bituminous deposit, which is likely linked to closed
paleogeography conditions. They suggested a water stratification
considering the oxygenated water and minimum oxygen layer
(minimum O2/O2) interface as a favorable zone for the phosphate
deposits and the minimum oxygen zone as the favorable zone for
the bituminous deposits. The phosphate sediments, including those
of the Foum Kheneg Formation studied herein, have then been
deposited at the beginning of the Maastrichtian and Paleogene
transgressive phases in an area where the oxidation-reduction
conditions are neither too oxidizing nor reducing.
In their study of the relationship between the atmospheric cir-
culation and the oceanic distribution of organic-rich rocks, Parrish
and Curtis (1982) have stated that the organic-rich rocks distribu-
tion is related to anoxic oceanic events, transgressions or to up-
wellings. And referring to the previous studies of the Middle Atlas,
we consider that the bituminous rocks are related to anoxic sea
conditions and the distribution of organic content in the phosphate
deposits is related to the MaastrichtianePaleogene transgression.
However, upwellings were not considered to be responsible for
phosphate deposits in the Middle Atlas.
5. Material and methods
5.1. The Bou Angueur section
The studied section (geographic coordinates: 33 337.9000 North
and 5 847.57” West) is located in the northern flank of the Bou
Angueur syncline, 48.61 km south of the Azrou city and close to
the national road N13 (Fig. 1B). The section is a part of the Foum
Kheneg Formation. It begins at its base with phosphate deposits
alternated with slightly phosphatic argillite beds (360 m thick)
(Figs. 2, 3). These phosphate levels are often fossiliferous and have
the following different aspects: silty-marly, clayey, marly-
calcareous, or sandy phosphates. They are surmounted by an
alternation of bioclastic limestone and marly beds (150 m thick).
The upper part (330 m thick) of the series is marked by in-
tercalations of phosphate levels with beds of marl, marly-clay, clay
or slightly phosphatic sand. These phosphates are characterized by
their sandy appearance and fossiliferous content (fish teeth,
biogenic debris ...). This section was previously assigned to the
Upper CretaceousePaleogene interval, based on lithological cor-
relations with other sections in the Middle Atlas (Herbig and
Fechner, 1994). This needed to be refined to stage level in order
to identify the K/Pg boundary, which is the main purpose of this
palynological study.
5.2. Methods
Twenty-seven samples were collected equidistantly every
30 cm in the Bou Anguer section and treated according to the
standard palynological procedure as described by Slimani et al.
(2016). Sample surfaces were carefully washed to remove con-
taminants, dried at 60  C and then crushed to facilitate the acid
attacks. 40e100 g of each sample were dissolved in cold hydro-
chloric acid HC1 (10%) for 1 h to remove the carbonate fraction,
followed by two successive attacks for 96 h in cold hydrofluoric acid
HF (40%) to eliminate the silicates. Neoformed silicofluorides were
removed by boiling the remaining material in HC1 (20%) for 20 min.
Due to the richness in amorphous organic matter, the residues were
treated with oxidizing reagent as nitric acid HNO3 (50%) and po-
tassium chlorate KClO3 (99%) for 45 min, followed by neutralization
in potassium hydroxide KOH (10%) to reveal the palynomorphs.
After each chemical treatment, the samples were washed with
distilled water until they are neutralized. Residues were sieved on a
nylon screen with a mesh size of 15 mm, stained with safranin
(C20H19ClN4), in order to enhance palynomorph colors, and then
mounted on microscope slides using glycerine jelly. Three to four
slides per sample were prepared. 400 organic-walled palyomorphs
(spores, pollen, acritarchs, dinoflagellate cysts) per sample were
counted where possible and the remaining palynomorphs in each
slide were checked for rare biostratigraphic dinocyst markers and
exceptionally well-preserved specimens for photographic
 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
Fig. 2. Lithostratigraphy, sampling points and biostratigraphic dinocyst events in the CretaceousePaleogene boundary section of Bou Angueur, Middle Atlas, Morocco. Succession of
sample numbers is from base to top of the section.
documentation. England Finder (EF) coordinates of the figured
specimens are given in Figs. (6e10) captions. The nomenclature of
the dinocyst taxa and references to their authors follow the data-
base DINOFLAJ3 (Williams et al., 2017). Slides were examined under
an Olympus BX53 transmitted light microscope and dinocyst
photomicrographs were taken using a digital Olympus C-400 Zoom
camera. All dinocyst taxa identified during this study are listed
alphabetically in Appendix A and their stratigraphic distribution is
plotted in Appendix B. All palynological slides are stored in the
botanical collection of the National Herbarium of Rabat (RAB),
Scientific Institute, Mohammed V University of Rabat, Morocco.
England Finder (EF) coordinates of figured specimens are given in
the figure captions.
For the quantitative analyses, the relative abundances of dino-
cyst groups, often considered as paleoenvironmental indicators,
were calculated and the results were expressed in percentages (%)
of the total count (Fig. 5, Appendix B). The following categories are
frequently used in the text: few (1e5%), rare (5e10%), common
(10e20%), abundant (20e40%) and superabundant (>40%). The
following ratios (cf. Versteegh, 1994; Guasti, 2005), frequently used
5
as paleoenvironmental proxies, were calculated using the number
(n) of counted palynomorphs (Appendix B):
1) the peridinioid/gonyaulacoid dinocyst ratios [P/G ¼ nP/
(nP þ nG)] are used to estimate the sea surface productivity;
2) the sporomorph (spores and pollen)/dinocyst (dinoflagellate
cysts and acritarchs) ratios [S/D ¼ nS/(nD þ nS)] are used to
evaluate the relative influences of terrestrial versus marine
organic material;
3) the inner neritic dinocyst/outer neritic to oceanic dinocyst ratios
[IN/ON ¼ nIN/(nIN þ ON)] are used to estimate the relative sea
level change;
4) the warm-water dinoflagellate/cold-water dinoflagellate ratios
[W/C ¼ nW/(nW/nW þ nC) are used to estimate changes in the
sea surface temperatures.
The carbonate analyses were achieved following the French
norm NFP 94-048 (AFNOR, 1996), which is based on the measure-
ment of carbon dioxide (CO2) volume evolved by the hydrochloric
acid (HCl) attack.
6 S. Chakir et al. / Cretaceous Research 106 (2020) 104219

Fig. 3. Field photographs of the K/Pg boundary section of Bou Angueur.

Fig. 4. Percentage of the main groups of palynomorphs (dinocysts, sporomorphs, acritarchs) across the CretaceousePaleogene boundary in the Bou Angueur section, Middle Atlas,
Morocco. (A) Maastrichtian, (B) Danian.
 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
Fig. 5. Quantitative results for paleoenvironmental and paleoclimatic reconstructions across CretaceousePaleogene boundary in the Bou Angueur section, Middle Atlas, Morocco. Horizontal bands reflect only warming and cooling
episodes inferred from the acmes of the thermophilic dinocyst taxa. The latest Maastrichtian cooling episode is not shaded, because it is not reflected by an acme but only by the consistence presence of the Isabelidinium group.

7
8 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
 S. Chakir et al. / Cretaceous Research 106 (2020) 104219 9

6. Results and discussion 6.1.2. Danian

6.1. Dinocyst stratigraphy and age assessment
The palynological analysis of the studied section required a
lithological description (see material and methods) and a high
resolution and equidistant (every 30 cm) sampling. The analyzed
samples contain well-preserved, rich and diverse dinoflagellate
cyst assemblages, reaching 80e99.5% of total palynomorphs
(Fig. 4). Other marine palynomorphs, such as acritarchs are rather
rare (up to 5.9%). The continental palynomorphs (pollen and
spores) are also rare (up to 8%), except for sample T22 where they
are common (16.66%) (Fig. 5). We identified a total of a hundred and
twenty-four dinoflagellate cyst species, among which thirty-two
species are shaded in grey in Appendix B, since they are consid-
ered as biostratigraphic markers of the upper Maastrichtian and
lower Danian.
Age assessment of this section is based on comparisons with
other studies of Maastrichtian to Danian dinocyst assemblages
from several sections in the Northern Hemisphere, mainly the
Mediterranean sections.
6.1.1. Upper Maastrichtian
From sample T1 (marly/silty phosphates) at the base of the
section upwards to sample T17 (yellowish marl), the recorded
dinocyst association can be assigned to the upper Maastrichtian,
based on the following dinocyst marker events (Appendix B): The
first occurences (FOs) of Cordosphaeridium inodes subsp. inodes,
Deflandrea galeata and Glaphyrocysta perforata. The FOs of these
species characterize the late Maastrichtian in the Northern Hemi-
sphere (Herngreen et al., 1986; Firth, 1987; Brinkhuis and
Zachriasse, 1988; Soncini and Rauscher, 1988; Dupuis et al., 2001;
Slimani, 1995, 2001; M’Hamdi et al., 2013, 2014; Slimani et al.,
2010, 2011, 2016; Fensome et al., 2008; Gue
de

et al., 2014). We
also used the FOs of other markers of the latest Maastrichtian, such
as Apteodinium fallax, Carpatella septata, Cerodinium medi-
terraneum, Cerodinium striatum, Conosphaeridium lifum, Glaphyr-
ocysta ordinata, Kenleyia leptocerata, Kenleyia lophophora,
Lanternosphaeridium cf. lanosum, Lejeunecysta hyalina, Lejeunecysta
izerzenensis, Phelodinium elongatum, Palaeocystodinium bulliforme,
Renidinium gracile and Exochosphaeridium sp. complex of Benson
(1976). They were recorded in the middle latitudes of the North-
ern Hemisphere, especially in the Mediterranean regions (Slimani
et al., 2008, 2010, 2012, 2016; Mohamed et al., 2012, 2013;
Slimani and Toufiq, 2013; Gue
de
et al., 2014; M’Hamdi et al.,
2013, 2015; Açıkalın et al., 2015; Soliman and Slimani, 2019). The
last occurrences (LOs) of Dinogymnium spp. were considered as
markers of the latest Maastrichtian (Williams et al., 2004), but they
were also observed in the early Danian by other authors, such as
Brinkhuis and Schiøler (1996) and Slimani et al. (2010). The LOs of
Dinogymnium acuminatum and Dinogymnium albertii are observed
in sample T17, while most of the other species of this genus cross
the CretaceousePaleogene boundary and disappear in the lower
Danian (Appendix B) (see more detail in the paragraph 6.1.3).
The basal Danian is recognized from sample T18 to sampleT27,
based on the FOs of Carpatella cornuta, Danea californica, Lanter-
nosphaeridium reinhardtii and Senoniasphaera inornata. The FOs of
these species mark the early Danian in northwestern Europe
(Hansen, 1977; Hultberg, 1985; Slimani, 1995, 2000, 2001; Slimani
et al., 2011) and United States of America (Habib et al., 1992, 1996;
Moshkovitz and Habib, 1993). They were also used to identify the
K/Pg boundary in Tethyan regions of Morocco (Slimani et al., 2010,
2016; Slimani and Toufiq, 2013; Gue
d e

et al., 2014), Tunisia
(Brinkhuis and Zachariasse, 1988; Dupuis et al., 2001; M'Hamdi
et al., 2013, 2015; Vellekoop et al., 2015) and Turkey (Açıkalın
et al., 2015). These dinocyst bioevents were considered as
worldwide markers of the Danian by Williams and Bujak (1985),
Haq et al. (1987) and Williams et al. (2004). In the Bou Angueur
section, the FO of Danea californica is recorded in sample T18,
while the FOs of Lanternosphaeridium reinhardtii and Seno-
niasphaera inornata are observed in sample T22. The occurrence
of S. inornata is restricted to the early and middle Danian
(66.00e62.60 Ma) (Williams et al., 2004). Carpatella cornuta first
occurs in sample T18 and shows a consistent presence from
sample T23 to the top of the section. Consequently, the strati-
graphic distribution of all these species suggests a Danian age for
the sample interval (T18eT27). Areoligera constricta, indicating
the Danian (Vieira et al., 2017), occurs in the sample T27, con-
firming the Danian age for this interval. Impagidinium maghri-
bensis, described for the first time in the earliest Danian in
Morocco (Slimani et al., 2008) and subsequently recorded in the
early Danian (Slimani et al., 2010, 2016; M'Hamdi et al., 2013,
2015; Gue
d e

et al., 2014; Mohamed et al., 2012), is recorded
herein in sample T18 and consequently supports the Danian age
suggested herein for this interval. The FO of Cassidium fragile,
previously reported at the MaastrichtianeDanian transition, just
below the FO of Danea californica (Drugg, 1967; M'Hamdi et al.,
2013) or in the basal Danian (Slimani et al., 2010, 2016; Gue
d e


et al., 2014; Açıkalın et al., 2015; Vellekoop et al., 2015) is also a
worldwide marker of the Danian. In the Bou Angueur section, the
FO of Cassidium fragile is observed in sample T16, below the FO of
Danea californica in sample T18.
All These findings from the Bou Angueur section allowed us to
assign the interval from sample T1 to sample T16 to the upper
Maastrichtian and the interval from sample T18 toT28 to the Dan-
ian, respectively. Therefore, the K/Pg boundary likely lies between
samples T17 and T18, most likely in the thin clayey layer located
between the yellowish clay and the phophatic clayey sand.
Geochemical analyses are necessary to check the presence or not of
an iridium anomaly in this interval.
6.1.3. Biostratigraphic discussion
The K/Pg boundary section in the Bou Angueur syncline diplays
a succession of regional and global biostratigraphic dinocyst events
(Fig. 2, Appendix B), including the FOs of Carpatella septata, Kenleyia
leptocerata, Kenleyia lophophora and Glaphyrocysta perforata, indi-
cating the uppermost Maastrichtian, followed by the FOs of

Fig. 6. Dinoflagellate cysts across the CretaceousePaleogene boundary in the Bou Angueur section, Middle Atlas, Morocco. Scale bar in Fig. A represents 40 mm for all specimens
(AeP). A. Elytrocysta druggii, sample T12, slide 1, EF Y45, focus on the archeopyle and processes. B. Impagidinium maghribensis, sample T18, slide 3, EF E27, dorsal surface, focus on
the precingular archeopyle. C. Palaeohystrichophora infusorioides, sample T5, slide 2, EF Y23/4, optical section showing an antapical horn and processes. D. Spiniferites membranaceus,
sample T21, slide 1, EF Z25/3, dorsal surface. E, F. Carpatella septata, sample T10, slide 1, E. Ventral surface, F. Dorsal surface showing the precingular archeopyle. G. Cassidium fragile,
sample T16, slide 1, focus on the apical archeopyle. H. Operculodinium centrocarpum, sample T15, Slide 2, EF Q30, dorsal surface showing the precingular archeopyle. I. Apteodinium
fallax, sample T28, slide 1, EF T34/1, dorsal surface. J. Kallosphaeridium yorubaense, sample T14, slide 2, EF W37, focus on the archeopyle and attached operculum. K, L. Danea
californica, K. Sample T29, slide 3, EF T37, dorsal surface showing the precingular archeopyle and processes, L. Sample T29, slide 3, EF V25, dorsal surface showing the precingular
archeopyle and processes. M. Kenleyia lophophora, sample T15, slide 1, EF D31, lateral surface. N. Cordosphaeridium fibrospinosum, sample T8, slide 1, EF W40, focus on the precingular
archeopyle and fibrous processes. O. Cordosphaeridium exilimurum, sample T10, slide2, EF Z39/1, dorsal surface. P. Cordosphaeridium inodes subsp. inodes, sample T 10, slide 2, EF Z27,
dorso-lateral surface.
10 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
Cassidium fragile, Danea californica, Carpatella cornuta and Lanter-
nosphaeridium reinhardtii indicating the lower Danian.
The dinocyst species Palynodinium grallator of which the LO is
considered as a marker of the latest Maastrichtian or the
MaastrichtianeDanian boundary in the northern higher-latitudes
(Hansen, 1977, 1979a, b; Schiøler and Wilson, 1993; Habib et al.,
1996; Schiøler et al., 1997; Slimani et al., 2011), is not found in
the Bou Agueur section. The previous records of this cold-water
species above the K/Pg boundary in the Tethyan regions was
interpreted as the result of a southward migration caused by a
cooling event in the earliest Danian (Brinkhuis et al., 1998; Slimani
et al., 2010, 2016; Vellekoop et al., 2015). The absence of this species
in the studied section might not only be related to the climatic
conditions (sea surface temperature), but also to its extreme scar-
city, unfavorable environmental conditions or oceanic paleocurrent
(see also Gue
de

et al., 2014). Nevertheless, another high-latitude
cold-water dinocyst species Isabelidinium cooksoniae whom the
LO is also known as a marker of the latest Maastrichtian in Northern
Europe (Schiøler and Wilson, 1993; Slimani et al., 2011), first occurs
at the top of the Maastrichtian with low abundances in the studied
section. But, it becomes abundant and even shows two abundance
spikes above the K/Pg boundary in sample T18 (11.3%) and T21
(21%). This may also be the result of a possible southward migration
caused by repeated pulses of cooling, mainly during the early
Danian.
The dinocyst species Manumiella seelandica, which is known by
its global abundance spike below the K/Pg boundary, related to the
latest Maastrichtian global cooling (Habib and Saeedi, 2007), is not
recorded in the studied section. This absence may be explained by a
hiatus of the uppermost part of the upper Maastrichtian sediments,
which may be due to syn-sedimentary vertical movements (see
also Mebrouk et al., 2009), lack of sampling or unfavorable envi-
ronmental conditions. In northern Morocco, the acme of Manu-
miella seelandica was observed in the uppermost Maastrichtian or
below the K/Pg boundary in probably more stable sedimentary
conditions in the “phosphate Plateau” (Rauscher and Doubinger,
1982; Rauscher, 1985; Soncini, 1992) and in the External Rif
(Slimani et al., 2010, 2016).
The LOs of Dinogymnium spp. and its morphologically related
Alisogymnium euclaense are, for certain authors, useful for identi-
fying the K/Pg boundary in the Northern Hemisphere (Williams
et al., 2004; Slimani et al., 2011, 2016; M'Hamdi et al., 2013; Guede

et al., 2014; Soliman and Slimani, 2019). However for others, these
taxa cross the K/Pg boundary to disappear later in the basal Danian
(Brinkhuis and Schiøler, 1996; Slimani et al., 2010; Açıkalın et al.,
2015; Vellekoop et al., 2015). This could be related to reworking or
also represents their final demise in the earliest Danian. In the Bou
Angueur section, Dinogymnium acuminatum and Dinogymnium
albertii last occur in the uppermost Maastrichtian (sample T17),
while Alisogymnium euclaense, Dinogymnium cretaceum and Dino-
gymnium westralium cross the K/Pg boundary and last occur within
the Danian in samples T22, T26, and T23, respectively.
6.2. Paleoenvironmental and paleoclimatic reconstructions
The paleoenvironmental reconstructions are inferred from
changes in the relative abundances of organic palyomorphs (spores,
Fig. 7. Dinoflagellate cysts across the CretaceousePaleogene boundary in the Bou Angueur section, Middle Atlas, Morocco. Scale bar in Fig. A represents 40 mm for Figs. AeK, scale
bar in Figs. J, L and M represent 40 mm. A, B. Areoligera constricta, sample T27, slide 10, EF Z48/3, A. Dorsal surface, B. Ventral surface. C. Areoligera senonensis, sample T14, slide 2, EF
O42/1, specimen in dorsal view. D, E. Glaphyrocysta perforata, sample T16, slide 1, D. Dorsal surface, E. Ventral surface. F. Areoligera coronata, sample T11, slide 2, EF J47, ventral
surface. G. Areoligera volata, sample T27, slide 10, EF Z54/2, dorsal surface. H, I. Renidinium gracile, sample T2, slide 10, H. Ventral surface with focus on the apical archeopyle, I. Dorsal
surface. J, K. Lanternosphaeridium reinhardtii, J. Sample T23, slide 1, EF X48, focus on the fibrous surface and on apical and antapical horns, K. sample T23, slide 1, EF V40/1, focus on
the precingular archeopyle. L. Exochosphaeridium sp. complexe of Benson (1976), sample T21, slide 1, EF T27, focus on the precingular archeopyle and distinctive apical process. M.
Pterodinium cingulatum subsp. reticulatum, sample T4, slide 1, EF L50, focus on the wall structure and sutural crests.
11
pollen, acritarchs and dinoflagellate cysts). This quantitative anal-
ysis is based on counting 400 palynomorphs in each slide. Amor-
phous organic matter (AOM) has not been taken into account, since
it was removed (refer back to chapter 5.2: methods) to extract
dinocysts, spores and pollen.
Although the dinoflagellate cysts do not represent the entire
planktonic community of motile dinoflagellates, they constitute an
excellent tool for paleoenvironmental reconstitutions. Changes in
the relative abundances of the gonyaulacoid and peridinoid cysts
reflect generally fluctuations in the depositional environment.
Harland (1973) noticed a relationship between open sea phases and
the increases of the gonyaulacoid cyst abundance and Davey (1979)
deduced a high abundance of peridinoid dinocysts in proximal
marine environments. Lister and Batten (1988) considered that the
relative abundances of the peridinoid and gonyaulacoid cysts could
be used to interpret the proximity to the coast and also the pale-
osalinity. An increase in the relative abundance of peridinoid cysts
indicates a low salinity, whereas the gonyaulacoid cysts indicate a
normal salinity.
The ecological interpretation of the dinocyst assemblages in this
study has been performed based on changes in the relative abun-
dances of selected dinocyst groups, which were considered in the
literature as environmental indicators and on P/G, IN/ON, S/D and
W/C ratios (see chapter 5.2: methods, for more detail) (Tables 1 and
2; Fig. 5). The selected dinocyst groups, followed by the included
dinocyst taxa between brackets, are: Kleithriasphaeridium (Flor-
entinea spp., K. truncatum), Spiniferites (Achomosphaera crassipellis,
Spiniferites spp.), Areoligera (Areoligera spp., Glaphyrocysta spp.),
Cordosphaeridium (Cordosphaeridium spp., Lanternosphaeridium
reinhardtii, Kenleiya spp., Danea californica), Isabelidinium (Isabeli-
diniun spp., Chatangiella spp.), Pterodinium (Pterodinium cingulatum
subsp. reticulatum), Operculodinium (Operculodinium spp.), Palae-
ocystodinium (Andalusiella spp., Palaeocystodinium spp.), Sene-
galinium (Senegalinium spp., Phelodinium spp., Lejeunecysta spp.,
Cerodinium spp., Deflandrea spp.). Seven paleoecological intervals
are described, based these parameters (Fig. 5):
6.2.1. Interval I (500 cm up to 410 cm below the K/Pg boundary)
Interval I (upper Maastrichtian) extends from sample T1 to
sample T4, collected in the greenish to blackish marly/silty phos-
phate (sample T1) at the base of the section and in the grayish
fossiliferous marly phosphate (samples T2eT4). The CaCO3 content
is moderate (40e50%). This interval shows a moderate dinocyst
diversity (20e30 species). The Senegalinium group (24e38.5%) is
slightly more abundant than the Spiniferites group (18.5e21%) in
samples T1 and T2. The Spiniferites group was considered as a
cosmopolitan, proliferating in neritic and oceanic environments
with a stable salinity and even more in high-productivity envi-
ronments and upwelling zones (Brinkhuis, 1994; Zonneveld et al.,
2013). In the other hand, the Senegalinium group proliferates in
shallower neritic environments with high productivity (Brinkhuis
and Zachariasse, 1988; Sluijs et al., 2009; Sluijs and Brinkhuis,
2009). These findings suggest intermediate inner-outer neritic
marine environments and moderate productivity conditions for the
two samples T1 and T2 covering the marly/silty phosphate. This is
supported by an intermediate relative sea level inferred from a
moderate IN/ON ratios (0.5e0.6) in the two samples. Samples T3
12 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
 S. Chakir et al. / Cretaceous Research 106 (2020) 104219 13

and T4 from the marly fossiliferous phosphate are characterized by
very low IN/ON ratios (0.1e0.5) and P/G ratios (0.1), indicating a
relative sea level rise and low productivity, respectively. This is
confirmed by a decrease in the relative abundance of the Sene-
galinium group (4.3e5.5%) and an increase of the Spiniferites group
(33.3e43%). From sample T3 to sample T4, the Areoligera group
records a decrease in its relative abundances (40.5%e0.5%) coupled
with an increase of the Pterodinium group (0%e40.8%). The abun-
dance of the Pterodinium group suggests a stable oceanic environ-
ment (Pross and Brinkhuis, 2005; Sluijs et al., 2005), whereas the
Areoligera group is associated to shallow littoral environments with
high energy. The high abundance Pterodinium group may indicate a
transgression and a sea level rise (Brinkhuis, 1994; Guasti et al.,
2005; Sluijs et al., 2005; Slimani et al., 2010). As a result, these
dinocyst groups reflect a littoral environment for sample T3 fol-
lowed by a short period of more offshore conditions related to very
likely important changing ocean currents or a short period of
transgression following an eventual collapse of the Bou Angueur
area caused by an extensive tectonic activity for sample T4.
6.2.2. Interval II (380 cm up to 290 cm below the K/Pg boundary)
Interval II corresponds to the sample interval T5eT8, covering
the calcareous-marly phosphate (sample T5) and the lower part of
the clayey to slightly phosphatic marl (samples T6eT8). In this in-
terval, the CaCO3 content shows a decrease from 60% in sample T5
to 20% in sample T8, related to the change from calcareous sedi-
ment to clayey sediment. This interval is characterized by a mod-
erate dinocyst diversity (20e40 species) and fluctuations in the
relative abundances of the cosmopolitan Spiniferites group
(16.3e31%) against the inner neritic Senegalinium group
(7.3e43.3%) and in the P/G (0.1e0.5) and IN/ON (0.4e0.8) ratios.
The high-energy coastal Areoligera group is abundant (24%) in
samples T5 and T8. All these results suggest an intermediate neritic
environment with slight fluctuations in abundance of outer and
inner neritic dinocyst groups, probably due to fluctuation in
nutrient availability.
6.2.3. Interval III (260 cm up to 170 cm below the K/Pg boundary)
Interval III covers the upper part of the clayey to slightly
phosphatic marls, extending from sample T9 to sample T12. The
CaCO3 content is lower (20e30%) in comparison with the rest of
the section. This interval is characterized by a high dinocyst di-
versity (up to 60 species) e the highest in the studied section e
and a dominance of the peridinoid cysts, including the Sene-
galinium group (10e30%) and the Palaeocystodinium group
(30e70%), the Spiniferites group is rare. The IN/ON (0.7e0.9) and
the P/G (0.4e0.7) ratios are high, reflecting a decrease of the
relative sea level and an increase of the productivity, respectively.
The Palaeocystodinium group, also called Andalusiella-Palae-
ocystodinium A-P complex (Brinkhuis et al., 1998; Vellekoop et al.,
2015), is the most dominant. It is composed of two tropical-
subtropical genera (Andalusiella and Palaeocystodinium), which
are associated with inner neritic environments typical of up-
welling systems and warm-water marine conditions (Wall et al.,
1977; Lentin and Williams, 1980; Brinkhuis et al., 1998;
Vellekoop et al., 2015). The high abundance of the
Palaeocystodinium group is reflected by high W/C ratios, reaching
the highest values in this interval (0.7e1). A similar interval with
high abundances of the Palaeocystodinium group was recorded by
Vellekoop et al. (2015) in the interval of 2e4 m below the K/Pg
boundary, within the Abathomphalus mayaroensis foraminifer
Zone (upper Maastrichtian) of the Elles section (Tunisia). In
summary, the interval III records a marginal inner neritic envi-
ronment, but under warm climate conditions inferred from high
abundances of the thermophilic Palaeocystodinium group. These
warm climatic conditions may be correlated with one of the
warming phases e recorded previously during the last 500 kyrs of
the Maastrichtian, prior to the K/Pg boundary e likely related to
Deccan volcanism (Olsson et al., 2002; Stüben et al., 2003; Keller
et al., 2012, 2018; Punekar et al., 2014).
6.2.4. Interval IV (140 cm below the K/Pg boundary)
Interval IV extends from sample T13 to sample T16. It covers
the limestone layer (sample T13), the greenish clayey marl
(samples T13eT15) and the limestone layer (sample T16). The
CaCO3 content (up to 85%) in the two limestone layers is the
highest in the section. The P/G (0.2e0.3) and the IN/ON (0.4e0.5)
ratios decrease, compared with the previous interval, suggesting a
decrease in the paleoproductivity and an increase in the relative
sea level, respectively. The cosmopolitan Spiniferites (18e40%) and
Operculodinium (10e20%) groups are dominant, while the
restricted inner neritic Senegalinium group records a significant
decrease in the relative abundance (<30%). The coastal Areoligera
group is also well presented, but shows a decrease from 38% in
sample T13 to 19% in sample 17. The warm-water Palae-
ocystodinium group has dropped from 70% to 3% and the W/C
ratios from 1 to 0.3. However, the Isabelidinium group, an indi-
cator of near-shore and cold-water environments (Dettman and
Thomson, 1987; Stover et al., 1996; Thorn et al., 2009; Castro
and Carvalho, 2015), shows consistent occurrences (7.5e8.5%).
These results may suggest an intermediate (inner-outer neritic)
environment with more offshore conditions, possibly due to a
decrease in nutrient availability, related to a decrease of upwell-
ing strength or freshwater input. The consistent first appearance
and the increase in abundance of the cold-water Isabelidinium
group during this interval may suggest a cooling, probably related
to a relatively cooling phase of the late Maastrichtian (Stüben
et al., 2003; Keller et al., 2012; Vellekoop et al., 2015). Accord-
ing to Vellekoop et al. (2015), this cooling reflects the aftermath of
a Late Cretaceous warming event, probably related to the Deccan
volcanism (Olsson et al., 2002). The cooling phase in this interval
(IV) may then reflect the aftermath of the warming phase inferred
from high abundances of the thermophilic Palaeocystodinium
group in the previous interval (III). The interval IV may coincide
with the pre-impact phase recognized in the Tethyan sections of
Ouled Haddou, El Kef and Okçular sections by Vellekoop et al.
(2017). The absence of the Manumiella seelandica spike, which is
related to global cooling in the latest Maastrichtian (Habib and
Saeedi, 2007; Slimani et al., 2010, 2016; Slimani and Toufiq,
2013; Vellekoop et al., 2015), may be explained by a hiatus of
the uppermost part of the uppermost Maastrichtian sediments,
lack of sampling or unfavorable environmental conditions.

Fig. 8. Dinoflagellate cysts across the CretaceousePaleogene boundary in the Bou Angueur section, Middle Atlas, Morocco. Scale bar in Fig. A represents 40 mm for all specimens (A
to M). A, B. Florentinia buspina, sample T3, slide 3, EF Z46, A. ventral surface with focus on the processes, B. focus on the archeopyle combining the precingular plate P (300 ) and apical
plates (10 e40 ). C. Carpatella cornuta, sample T25, slide 2, EF Z38, focus on the wall structure and apical and antapical horns. D, E. Kleithriasphaeridium truncatum, sample T6, slide 1, EF
P32, D. focus on the precingular archeopyle, E. focus on the ventral surface. F. Alisogymnium euclaense, sample T5, slide 2, EF U37/4. G. Dinogymnium cretaceum, sample T10, slide 2,
EF Z39/3. H. Pierceites pentagonus, sample T21, slide 1, EF O47/3, focus on the intercalary archeopyle 3I. I. Chatangiella manumii, sample T6, slide 1, EF W29/1, dorsal surface. J.
Dinogymnium acuminatum, sample T14, slide 1, EF O39/3. K. Isabelidinium cooksoniae, sample T15, slide 1, EF G50/1, dorsal surface with focus on the granulose surface and the
intercalary archeopyle. L. Chatangiella sp. A of Schiøler and Wilson (1993); sample T21, slide 2, EF L40, focus on granulose surface and cingular spines. M. Isabelidinium bakeri, sample
T14, slide 1, EF V49/1, dorsal surface with focus on the intercalary archeopyle.
14 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
6.2.5. Interval V (30 cm around the K/Pg boundary)
Interval V covers the yellowish marl below the presumed K/Pg
boundary (sample T17) and the first 10 cm thick of the sandy
phosphate (sample T18) above the K//Pg boundary (thin clayey
layer). The CaCO3 content decreases abruptly from 80% in the
previous interval to 30% in this interval. This decrease may be
related to a sudden reduction of primary productivity at the K/Pg
boundary due to the disappearance of most of Cretaceous calcar-
eous organisms (Keller and Lindinger, 1989). The dinocyst biodi-
versity is moderate (30e40 species). In the yellowish marl (sample
T17), the P/G (0.5) and IN/ON (0.5) ratios are moderate and in
agreement with moderate relative abundances of the gonyaulacoid
cysts and peridinoid cysts, reaching 54.3% and 45.9%, respectively.
This means that the percentage of the peridinoids increases due to
the increase of the cold-water Isabelidinium group, suggesting more
inner neritic environment in comparison with the previous inter-
val. In the sandy phosphate (sample T18), ~10 cm above the KPg
boundary, the gonyaulacoid groups (Spiniferites, Areoligera, Oper-
culodinium) are rare to absent, whereas the peridinoid groups
(Senegalinium, Isabelidinium) become superabundant (>80%) due to
the sudden increase (up to 40%) in the relative abundance of the
cold-water taxa, including the high-latitude cold-water Isabelidi-
nium cooksoniae and Chatangiella sp. A of Shiøler and Wilson
(1993). The W/C ratio remains below 0.5, suggesting cold-water
conditions. Similar records of this cold-water dinocyst incursion,
expressed by other typical higher-latitude dinocyst taxa (Paly-
nodinium grallator, Cribroperidinium spp.), have been reported at
the base of the planktonic foraminifer P0 Zone at El Kef (Brinkhuis
et al., 1998), Ouled Haddou (Slimani et al., 2010; Slimani and Toufiq,
2013) and Elles (Vellekoop et al., 2015). The significant increase in
relative abundance of the higher-latitude Isabelidinium group may
be correlative with end of the first cooling pulse, interpreted by
Vellekoop et al. (2014, 2015) as being related to an “impact winter”
in the first months to decades following the Chixculub impact.
Geochemical analyzes in the yellowish clay and the first centime-
ters of the sandy phosphate (between sample T17 and sample T18),
comprising the K/Pg boundary are needed in order to verify the
presence of an iridium anomaly and reveal the rapid cooling event
following the Chixculub impact in the studied section. The IN/ON
ratios reach their maximum value (1), the highest in this section. All
these findings suggest a marginal inner neritic marine environment
for the yellowish marl (sample T17) and the sandy phosphate
(sample T18), with a strong sea-level decrease. The increase of the
percentage of sporomorphs (up to 10%) compared to total paly-
nomorphs through the section and the increase of the S/D ratio (up
to 0.15) corroborate the previous suggestion. It should be noted that
the slight increase in sporomophs in the earliest Paleocene of the
Bou Angueur section is very similar to the record of Elles (Vellekoop
et al., 2015).
6.2.6. Interval VI (40e130 cm above the K/Pg boundary)
This interval consists of dark clayey marl (sample T19) and
slightly phosphatic clayey sand (samples T20eT22). The CaCO3
content is still low (less than 20%) in the basal Danian. The dinocyst
biodiversity varies from 20 to 40 species per sample. The clayey
marl (sample T19) is characterized by a dominance of the Sene-
galinium group (70%) and a high IN/ON ratio (0.8), indicating fresh-
Fig. 9. Dinoflagellate cysts across the CretaceousePaleogene boundary in the Bou Angueur section, Middle Atlas, Morocco. Scale bar in Fig. A represents 40 mm for all specimens
(AeM). A. Cerodinium speciosum, sample T10, slide 1, EF K53/4, dorsal surface with focus on the intercalary archeopyle. B. Senegalinium microgranulatum, sample T5, slide2, EF K30,
focus on the granulose endocyst. C. Senegalinium obscurum, sample T27, slide 10, EF Q41/2, focus on the intercalary archeopyle. D. Senegalinium laevigatum, sample T10, slide 2, EF
Z40/4, focus on the intercalary archeopyle and operculum. E. Lejeunecysta hyalina, sample T15, slide 1, EF K34. F. Deflandrea galeata, sample T3, slide 1, EF G48, dorsal surface. G.
Senegalinium bicavatum, sample T14, slide 1, EF X49. H. Lejeunecysta izerzenensis, sample T10, slide 3, EF H36/2, focus on the denticulate cingular crests. I. Phelodinium magnificum,
sample T7, slide 1, EF W30/4, dorsal surface with focus on the intercalary archeopyle. J. Lejeunecysta globosa, sample T3, slide 1, EF Q44/4. K. Phelodinium elongatum, sample T14, slide
2, EF G50/1.
15
water and high nutrient conditions (Sluijs and Brinkhuis, 2009).
This bloom may coincide with the disaster phase recognized in the
Tethyan sections by Vellekoop et al. (2017). A similar strong in-
crease in the abundance of the Senegalinium group was recorded
within the foraminifer Guembelitria cretacea Zone at Ouled Haddou
in Morocco (Slimani et al., 2010; Slimani and Toufiq, 2013). This
event was also observed, as an increase of the hexaperidinoids
(equivalent to our Senegalinium group), within the foraminifer P0
Zone and the lower part of P1 Zone at El Kef and Elles in Tunisia
(Brinkhuis et al., 1998; Vellekoop et al., 2015) and at Okçular in
Turkey (Vellekoop et al., 2017). According to Vellekoop et al. (2017),
this bloom of hexaperidinoids suggests that the earliest Paleocene
shelves of the Tethys were characterized by an increase in nutrient
availability in the photic zone due to collapse of the biological
pump following the mass extinction at the CretaceousePaleogene
boundary.
The phosphatic clayey sand (sample T20) is characterized by a
drop to 0% in the relative abundance of the Senegalinium group
coupled with a sudden increase in the relative abundance of the
opportunistic heterotrophic Palaeocystodinium group (60%), asso-
ciated with a sudden increase of the W/C ratio (0.9). A similar spike
of the Palaeocystodinium group has been recorded, as a bloom of the
Andalusiella-Palaeocystodinium A-P complex, at the top of the
planktonic foraminifer P0 Zone in the Tethyan regions. The spike
was correlated with the episode of global warming following the K/
Pg boundary as a result of the release of greenhouse gases into the
atmosphere (Kring, 2007; Vellekoop et al., 2014, 2015, 2018). In
contrast to sample T20 that is highly dominated by the peridinoid
dinocysts (Palaeocystodinium group), sample T 21 e also from the
phosphatic clayey sand e records a high abundance of gonyaula-
coid dinocysts, represented mainly by the cosmopolitan Spiniferites
group (38%), and moderate abundance (20%) of the cold-water
Isabelidinium group (mainly Isabelidinium cooksoniae). Sample T21
records also a decrease to 0.4 of the IN/ON ratio and a fall to 0.3 of
the W/C ratio, indicating an open outer neritic environment. As for
sample T22, collected at the top of the phosphatic clayey sand, the
dinoflagellate assemblage is characterized by a dominance (60%) of
the cold-water Isabelidinium group, among which Isabeldinium sp.
representing 19% of total palynomorphs. Isabelidinium sp. is
considered to be reworked because all specimens show signs of
transport (strong deformation and fragmentation), unlike other
dinocysts in the BouAngueur syncline succession. Reworking in this
sample can be supported by the presence, with a higher abundance
(3% of total palynomorphs) in the studied section, of the reworked
Upper Cretaceous Palaeohystrichphora infusorioides, coupled with a
higher abundance (20% of total palynomorphs) of spromorphs,
probably related to an increase input of eroded and also terrigenous
material. The high abundance of the Isabelidinium group is coupled
with a decrease to 0.2 of the W/C ratio. Similar records of typical
high-latitude dinocyst taxa have been reported in the planktonic
foraminifer Zone P1a at the El Kef stratotype section (Brinkhuis
et al., 1998) and Elles section (Vellekoop et al., 2015) and inter-
preted as related to a second cooling pulse following the initial
cooling pulse at the K/Pg boundary (Brinkhuis et al., 1998;
Vellekoop et al., 2014). Other subsequent cooling pulses were
recorded in both Tunisian sections, suggesting a kind of reverber-
ation of the initial cooling pulse at the K/Pg boundary, according to
16 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
 S. Chakir et al. / Cretaceous Research 106 (2020) 104219 17

Table 1
Paleoenvironmental interpretations of selected dinoflagellate cyst groups, after references.

Group Included taxa Paleoecological interpretation References

Areoligera Areoligera spp. Glaphyrocysta Nearshore shallow marine and high energy
spp. environments. Abundant occurrences are often
indicating a transgressive system.
Cordosphaeridium Cordosphaeridium spp. Open marine, inner neritic environment with a
L. reinhardtii Kenleiya spp. transgressive phase.
Danea californica
Operculodinium Operculodinium spp. Inner neritic to oceanic open marine environments.
cosmopolitan group. Acmes generally observed in
the upper shelf due to transportation.
Kleithriasphaeridium Florentinea spp. Deeper shelf waters.
K. truncatum
Pterodinium P.cingulatum subsp. reticulatum Oceanic open marine environment characterized by
low tolerance to nutrients and variations in salinity.
Its high abundance indicates an increase in sea level.
It's an autochthonous group (not transported from
shelf, but living in the oligotrophic pelagic water) in
the turbiditic sediments.
Spiniferites A. crassipellis Spiniferites spp. Several environment settings, such as oceanic to
restricted marine environments. This group can
become relatively cosmopolitan. Its highest
abundance was generally observed in the regions of
high productivity (upwelling areas and
environment influenced by river flow). This group
may represent a relative transgression following a
short term sea level fall.
Andalusiella Andalusiella spp. Tropical-subtropical, nearshore environment with a
Palaeocystodinium spp. rich nutrient availability. This group tolerates and
thrive in high productivity upwelling systems.
Isabelidinium Isabelidiniun spp. Chatangiella Inner neritic environment relatively close to the
spp. coastline, with considerable terrigenous influence.
Senegalinium Senegalinium spp. Phelodinium Shallow water with high-productivity level and
spp. Lejeunecysta spp. elevated nutrients, and influx of fresh water in
Cerodinium spp. Deflandrea spp. estuarine environments.
(Downie et al., 1971; 1994; Brinkhuis and
Zachariasse, 1988; Eshet et al., 1992; Firth, 1993;
Stover et al., 1996; Guasti et al., 2005; Sluijs et al.,
2008; Slimani et al., 2010; Chekar et al., 2018).
(Brinkhuis, 1994; Powell et al., 1996; Williams,
2006; Iakovleva, 2011; Oboh-Ikuenobe et al., 2017).
(Brinkhuis and Biffi, 1993; Brinkhuis, 1994; Stover
et al., 1996; Torricelli et al., 2006; Zonneveld et al.,
2013; Awad and Oboh-Ikuenobe, 2016).
(Leereveld, 1995; Prauss, 2001; Skupien, 2013).
(Wall et al., 1977; Brinkhuis and Zachariasse, 1988;
Marshall and Batten, 1988; Smelror and Leereveld,
1989; Dale, 1996; Prince et al., 1999, 2008; Harris
and Tocher, 2003; Crouch and Brinkhuis, 2005;
Skupien, 2011).
(Schrank, 1988; Brinkhuis and Zachariasse, 1988;
Zonneveld et al., 2013; Eshet et al., 1992; Brinkhuis,
1994; Brinkhuis and Schiøler (1996); Stover et al.,
1996; Brinkhuis et al., 1998; Pross, 2001; Sluijs et al.,
2003; Guasti et al., 2005; Sluijs et al., 2005; Torricelli
et al., 2006; Slimani et al., 2010; Iakovleva, 2015).
(Wall et al., 1977; Lentin and Williams, 1980;
Brinkhuis et al., 1998; Kring, 2007; Vellekoop et al.,
2014).
(Dettman and Thomson, 1987; Keating, 1992; Thorn
et al., 2009; Arai and Viviers, 2013; Castro and
Carvalho, 2015).
(Brinkhuis and Zachariasse, 1988; Eshet et al., 1992;
Firth, 1993; Nøhr Hansen and Dam, 1997; Gedl,
2004; Guasti et al., 2005; Sluijs, 2005;; Sluijs and
Brinkhuis, 2009; Iakovleva, 2015).

Vellekoop et al. (2015, p. 225). These results suggest a littoral inner This interval may encompass the disaster phase (sample T19)
neritic environment with cold-water conditions and a decrease in and the recovery phase (samples T20eT22) of Vellekoop et al.
the relative sea level associated to an important reworking for (2017).
sample T22.
In Summary, the interval VI records the succession of the
6.2.7. Interval VII (160e310 cm above the K/Pg boundary)
following events in ascending order:
Interval VII covers a succession of marly clay (samples
T23eT24), clayey sand (samples T25eT26) and clayey marl at the
1) an inner neritic environment with fresh-water and high
top of the section (sample T27). The CaCO3 content gradually in-
nutrient conditions reflected by the acme of the Senegalinium
creases to 50%. The dinocyst biodiversity is low (around 20 species).
group in the dark clayey marl (sample T19);
The gonyaulacoid dinocysts are highly dominant (96.9e100%),
2) an inner neritic environment with warm-water conditions re-
suggesting an open marine environment, whereas the peridinoid
flected by the acme of the opportunistic heterotrophic Palae-
dinocysts display an abrupt decrease in abundance (to 0%). The IN/
ocystodinium group, coupled with a fall in the relative
ON ratios are low (0.2) in sample T23 and moderate (around 0.6) in
abundance (close to 0%) of the Senegalinium group at the base of
the other samples. The P/G ratios are very low (0.03 in sample T23)
the phosphatic clayey sand (sample T20);
to null in the other samples. From base to top of this interval, the
3) a neritic environment with offshore conditions inferred from a
peridinoid dinocysts including the Senegalinium group are absent.
high abundance of the cosmopolitan groups, mainly the Spini-
However, the gonyaulcoid groups show gradual changes in the
ferites group for sample 21;
relative abundance. It displays a dominance of the cosmopolitan
4) an inner neritic environment with cold-water conditions, re-
Spiniferites group (60e100%) in the marly clay (samples T23 and
flected by the acme of the Isabelidinium group at the top of the
T24), an increase of the abundance of the Cordosphaeridium group
phosphatic clayey sand (sample T22).
(mainly Lanternosphaeridium reinhardtii (40e50%)) in the clayey

Fig. 10. Dinoflagellate cysts across the CretaceousePaleogene boundary in the Bou Angueur section, Middle Atlas, Morocco. Scale bar in Fig. A and H represents 40 mm, scale bar in
Fig. B represents 40 mm for Figs. BeD, and scale bar in Fig. E represents 40 mm for Figs. (EeG) and Figs. (IeL). A. Palaeocystodinium sp. cf. P. bulliforme, sample T1, slide2, EF J40/1, focus
on the intercalary archeopyle and operculum. B. Andalusiella inflata, sample T3, slide2, EF U43/3, focus on the wall structure. C. Andalusiella mauthei subsp. aegyptiaca, sample T23,
slide2, EF J34/4. D. Andalusiella polymorpha, sample T2, slide 10, EF W50/1, focus on the intercalary archeopyle. E. Palaeocystodinium benjaminii, sample T10, slide 1, EF U41/4, focus on
the apical and antapical horns. F. Palaeocystodinium australinum, sample T10, slide 2, EF Y35, specimen in dorsal view. G. Palaeocystodinium lidiae, sample T10, slide 1, EF Y52. H.
Palaeocystodinium golzowense, sample T10, slide 1, EF Z47/2. I. Andalusiella gabonensis, sample T10, slide 1, EF X55. J. Andalusiella mauthei subsp. punctata, sample T10, slide 1, EF U46/
2. K. Palaeocystodinium bulliforme, sample T5, slide 1, EF Z29/4. L. Andalusiella ivoirensis, sample T11, slide 2, EF T49, focus the sutural crests.
18 S. Chakir et al. / Cretaceous Research 106 (2020) 104219

Table 2
Latitudinal preferences of selected taxa in the latest Maastrichtian to earliest Danian interval, taken from the literature.

Low latitude preference References High latitude References

preference

Andalusiella spp.
Cerodinium pannuceum
Cerodinium striatum
Lejeunecysta spp.
Palaeocystodinium spp.
Phelodinium spp.
Senegalinium spp.
Areoligera volata
Cassidium fragile
Cerodinium mediterraneum
Conosphaeridium lifum
Lanternosphaeridium cf. lanosum
Danea californica
Kenleyia spp.
Lanternosphaeridium reinhardtii
Lingulodinium machaerophorum
Pierceites pentagonus
Trithyrodinium evittii
Kallosphaeridium yorubaense
Turbiosphaera galatea
(Malloy, 1972; Jain and Millepied, 1973; Riegel,
1974; Herngreen, 1975; Boltenhagen, 1977;
Lentin and Williams, 1980; Rauscher and
Doubinger, 1982; Schrank, 1987; Brinkhuis and
Zachariasse, 1988; Jan du Che ^ne, 1988; Oloto,
1989; Soncini, 1990; Eshet et al., 1992; Hoek
et al., 1996; Masure et, 1996; Masure et al.,
1998; El Beialy and Ibrahim, 1997; Brinkhuis
et al., 1998; Oboh-Ikuenobe et al., 1998; Ypes,
2001; Slimani et al., 2010, 2016; M'Hamdi et al.,
2013, 2015; Gue
de

et al., 2014; Slimani and
Toufiq, 2013; Vellekoop et al., 2015; Sẚnchez-
Pellicer et al., 1917; Gue
de
et al., 2019; Guler
et al., 2019; Soliman and Slimani, 2019).
(Drugg, 1967; Brinkhuis et al., 1998; Gue
de

Carpatella septata
et al., 2014; Slimani et al., 2016).
(Drugg, 1967; Brinkhuis et al., 1988; Slimani
et al., 2010, 2016; M'Hamdi et al., 2013; Gue
et al., 2014; Açıkalın et al., 2015; Vellekoop
et al., 2015).
(Oboh-Ikuenobe et al., 1998; Slimani et al.,
2008, 2010, 2016; Slimani and Toufiq, 2013;
M'Hamdi et al., 2013, 2014; Açıkalın et al.,
2015).
(Slimani et al., 2012, 2016; M'Hamdi et al.,
2015).
(Pre
v o
^ t et al., 1979; Rauscher and Doubinger,
1982; Slimani et al., 2010, 2016; Oboh-
Ekuenobe, 2017; Soliman and Slimani, 2019)
(Drugg, 1967; Erkmen et al., 1981; Brinkhuis
and Zachariasse, 1988; Brinkhuis et al., 1998;
Slimani et al., 2010, 2016; M'Hamdi et al., 2013;
Gue
de
et al., 2014; Açıkalın et al., 2015;
Vellekoop et al., 2015; Soliman and Slimani,
2019).
(Brinkhuis and Zachariasse, 1988; Brinkhuis
et al., 1998; Slimani et al., 2010, 2016;
Vellekoop et al., 2015; Oboh-Ekuenobe, 2017;
Soliman and Slimani, 2019).
(Moshkovitz and Habib, 1993; Habib et al.,
1996; Slimani et al., 2010; Açıkalın et al., 2015;
M'Hamdi et al., 2015; Vellekoop et al., 2015;
Soliman and Slimani, 2019).
(Brinkhuis, 1994; Londeix et al., 1999; Torricelli
et al., 2001; Zonneveld et al., 2001; Brinkhuis
et al., 2003; Sluijs et al., 2003).
(Rauscher and Doubinger, 1982; Soncini and
Rauscher, 1988; Soncini, 1990; Slimani et al.,
2010, 2016; Soliman and Slimani, 2019).
(Rauscher and Doubinger, 1982; Brinkhuis et al.,
1998; Yepes, 2001; Slimani et al., 2010, 2016;
Vellekoop et al., 2015; Guler et al., 2019;
Soliman and Slimani, 2019).
(Jan du Che^ne and Adediran, 1985; Soncini,
1990; Slimani et al., 2010, 2016; Gue
de

et al.,
d2014, 2019; Masure et al., 1998; Oboh-
Ekuenobe, 2017; Soliman and Slimani, 2019).
(El Beialy,1988, 1990; Slimani et al., 2010;
Soliman and Slimani, 2019).
Chatangiella spp. (Manum and Cookson, 1964; Vozzhennikova,
1967; Harland, 1973; McIntyre, 1975;
Iaonnides, 1986; Slimani, 2000, 2001; Fensome
et al., 2009; Slimani et al., 2011; Nohr-Hansen,
2012; Surlyk et al., 2013; McLachlan et al.,
2018).
(Willumsen, 2004, 2011).
Carpatella cornuta (Hansen, 1977; Hultberg, 1985; Damassa, 1988;

de

Brinkhuis et al., 1998; Slimani, 2000, 2001;
Slimani et al., 2011).
Areoligera senonensis (Brinkhuis et al., 1998; Slimani, 2000, 2001;
Cerodinium diebelii Slimani et al., 2011; Vellekoop et al., 2015).
Cerodinium glabrum (Gocht, 1969; Heilmann-Clausen, 1985;
Fensome et al., 2009).
Deflandrea galeata (Hansen, 1979a, b; Wilson, 1984,1987; Nøhr-
Hansen, 1994; Habib et al., 1996; Slimani, 2000,
2001; Slimani et al., 2011).
Cerodinium speciosum (Wilson, 1974; Schumacker-Lambry, 1977;
Herngreen et al., 1986; Marheinecke, 1992;
Schiøler and Wilson, 1993; Slimani, 1995, 2000,
2001; Schiøler et al., 1997; Brinkhuis et al.,
1998; Vellekoop et al., 2015).
Deflandrea denticulata (Wilson, 1971; Heilmann-Clausen, 1985;
Wilson et al., 1989; Berggren, 1994; Iakovleva
et al., 2000).
Exochosphaeridium sp. Benson (1976).
complexe of Benson
(1976)
Glaphyrocysta perforata (Hultberg, 1985; Brinkhuis et al., 1998; Slimani
et al., 2011; Vellekoop et al., 2015).
Hystrichosphaeridium (Wilson, 1974; Brinkhuis et al., 1998; Slimani,
tubiferum 2000, 2001; Slimani et al., 2011; Vellekoop
et al., 2015).
Isabelidinium spp. (Dettman and Thomson, 1987; Mao and Mohr,
1992; Schiøler and Wilson, 1993; Stover et al.,
1996; Schiøler et al., 1997; Slimani, 2000, 2001;
Thorn et al., 2009; Slimani et al., 2011; Castro
and Carvalho, 2015; Guler et al., 2019).

sand (samples T25 and T26) and an increase of the abundance of
the Areoligera group (60%) (mainly Areoligera senonensis) in the
clayey marl (sample T27), at the top of the section. Consequently,
this interval records open marine conditions during a transgressive
regime, suggesting more stable environmental conditions than
those recorded in the two previous intervals. The gradual increase
in the calcium carbonate content to ~50%, the highest in the sedi-
ments above the K/Pg boundary, may confirm a return to stable
marine conditions conducive to the increase of the productivity in
calcareous organisms as calcareous foraminifera and nannoplank-
ton. This interval may be related to the early Paleocene phase
recognized in the Tethyan sections by Vellekoop et al. (2017). The
abundance of the lower-latitude Lanternosphaeridium reinhardtii
(up to 47%), coupled with high W/C ratios (~0.8) may suggest
relatively warm-water conditions for samples T23 to and T26.
Whereas, the abundance of the high-latitude Areoligera senonensis
(40%), coupled with falls of the Cordosphaeridium group (0.1%) and
W/C ratio (0.1) in sample T27, may suggest relatively cold-water
 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
conditions that may be related to a cool water pulse, which is less
pronounced than the two previous presumed post ‘impact winter’
cooling water pulses. A. senonensis was considered as being a
higher-latitude species by Brinkhuis et al. (1998). These findings are
correlative with similar dinocyst records, reported within the
planktonic foraminifer P1 Zone in the Tethyan sections from
Morocco (Slimani et al., 2010; Slimani and Toufiq, 2013), Tunisia
(Brinkhuis et al., 1998; Vellekoop, 2014) and Turkey (Acikalin et al.,
2015; Vellekoop et al., 2017).
6.2.8. Paleoenvironmental discussion
In the Middle Atlas, the deposition of the phosphate succession
and the distribution of the organic content are related to the
MaastrichtianePaleogene transgression (see chapter 4 in this pa-
per). Upwellings have not been mentioned, as commonly known, as
being responsible of the deposition of the phosphate succession. It
should also be noted that the Middle Atlas was not directly con-
nected to the Atlantic Ocean, but through the “phosphate Plateau”
(Frizon de Lamotte et al., 2008) (see chapter 2 in this paper).
However, an important continental influence in the phosphate
deposits was reported, based on petrographic and mineralogical
analyzes by Mouflih et al. (2006). The quantitative changes of the
peridinoid dinocyst groups, especially the Senegalinium group may
then be more related to changes in continental nutrients avail-
ability and fresh-water conditions than to upwelling nutrients. On
the other hand, the subsequent cooling pulses recorded at Elles
(Tunisia) by Vellekoop et al. (2015) after the initial cooling pulse at
the K/Pg boundary were interpreted as resulted in a ‘reservoir’ of
cold water, that slowly dissipated to the surface ocean by upwell-
ing. In the Bou Angueur section, we also interpreted the second and
sole acme of the cold-water Isabelidinium group, after the initial
Isabelidinium acme following the K/Pg boundary, as related to a
second cooling pulse. In this area, this second cooling pulse may be
interpreted as a result of the dissipation of the deep cold water to
the surface ocean by the Paleogene transgression (at its beginning).
The indices of reworking and transport, indicated by Isabelidinium
sp. and sporomorphs, are in agreement with this transgression in
the uppermost part of the Bou Angueur section (see also paragraph
6.2.7: Interval VII). This transgression is confirmed by a sudden
increase of the gonyaulacoid dinocysts (mainly the cosmopolitan
Spiniferites group) and a sudden decrease of the inner neritic per-
idinoid dinocysts, recorded just above the Isabelidinium acme (in-
terval VII).
7. Comparison with other K/Pg sections
The dinoflagellate cyst assemblages recorded in the K/Pg
boundary interval of the Bou Angueur syncline are similar to those
from Tethyan sections. The most used KePg sections for compari-
son in this study are from Morocco (Slimani et al., 2010; Slimani and
Toufiq, 2013; Gue
de

et al., 2014; Slimani et al., 2016) and Tunisia
(Brinkhuis and Zachariasse, 1988; Brinkhuis and Leereveld, 1988;
Brinkhuis et al., 1998; Dupuis et al., 2001; M'Hamdi et al., 2013,
2014, 2015; Vellekoop et al., 2015). Other assemblages from Egypt
(Soliman and Slimani, 2019), Spain (De Conink and Smit, 1982;
Brinkhuis et al., 1998) and Turkey (Açıkalın et al., 2015;
Vellekoop, 2017) are also important for comparison. Most of rele-
vant dinocyst species markers and biostratigraphic events (first and
last occurrences, acmes), reported previously in the Tethyan sec-
tions are recorded in our section. However, the global spike of
Manumiella seelandica indicator of a global cooling below the K/Pg
boundary, Disphaerogena carposphaeropsis and the typical high-
latitude Palynodinium grallator are not found in the analyzed
samples. These differences might be related to differences in
paleogeographic and paleoceanographic settings and also
19
sedimentation rates and syn-sedimentary tectonic movements. The
multitude lithological changes in the phosphatic succession plus
the mostly inner to middle neritic marine environment inferred
from the changes in the relative abundances of selected dinocyst
groups through the Bou Angueur section (Fig. 5) suggest a depo-
sition in probably more proximal shelf setting than the Ouled
Hadou and El Kef sections. Nevertheless, the dinocyst records
suggest a relatively complete K/Pg boundary transition, covering
the most known global events following the K/Pg boundary Chic-
xulub impact. The studied section records similar events inferred
from the ecological responses of dinocysts to the K/Pg boundary
Chicxulub impact in the Tethtyan K/Pg sections, such as Ouled
Haddou (Slimani et al., 2010; Slimani and Toufiq, 2013), El Kef and
Okçular (Vellekoop et al., 2017). These events include the latest
Maastrichtian cooling, the brief period of global cooling or “impact
winter” following the K/Pg boundary Chicxulub impact, the global
warming episode following the impact winter and a second cooling
pulse. The ecological intervals IV, V, VI and VII, inferred from the
quantitative analysis in this study, are correlative, respectively with
the Maastrichtian or pre-impact phase, the disaster phase, the re-
covery phase and the early Paleocene phase, recently recognized
across the K/Pg boundary of Tethyan sections, including the Ouled
Haddou, El Kef and Okçular sections (Vellekoop et al., 2017).
However, the thickness of the sediment covering these events in
our section may be different, compared to those covering the
paleoenvironmental phases recognized by Vellekoop et al. (2017).
This may be due to sedimentation rates during these phases. The
disaster phase, which is characterized by strong abundances of
hexaperidinoids (equivalent to our Senegalinium group) within the
planktonic foraminifer P0 Zone (Vellekoop et al., 2017), covers the
first ~90 cm above the K/Pg boundary at El Kef (Brinkhuis et al.,
1998; Vellekoop, 2017). But it covers only ~30 cm above the K/Pg
boundary at Ouled Haddou (Slimani et al., 2010), Okçular
(Vellekoop et al., 2016) and Bou Angueur (base of interval VI). Also,
the recovery phase, which is recognized by a decrease in abundance
of the hexaperidinoids within the planktonic foraminifer Pa and
P1a zones (Vellekoop et al., 2017), encompasses ~530 cm at El Kef,
but only ~250 cm at Okçular and about ~100 cm at Bou Angueur
(interval VI). Therefore, the disaster phase in the studied section
records probably the same sedimentation rate as in the Okçular
section, while, it is nearly 3 times more condensed than in the El Kef
section. However, its recovery phase is 2.5 times more condensed
than in the Okçular section and 5 times more condensed than in the
El Kef section. The recovery phase in the Ouled Haddou section is
not taken into account in this correlation, since its upper limit is not
actually well constrained.
8. Conclusions
The phosphatic succession from the northern flank of the Bou
Angueur syncline near Azrou in the Middle Atlas, Morocco, yielded
rich and well-preserved palynological assemblages, dominated by
dinoflagellate cysts. This content allowed a new dating of the sec-
tion based on the first and last occurrences of dinocyst marker taxa,
and paleoenvironmental and paleoclimatic reconstructions infer-
red from quantitative analyses of carbonates and palynomorphs.
Dated for the first time, the section is revealed to encompass a
relatively complete record of the K/Pg boundary transition, suitable
for further geochemical analyses and other micropaleontological
studies.
We assigned the lower part of the section to the upper Maas-
trichtian and its upper part to the Danian, and identified the K/Pg
boundary between sample T17 and sample T18, most likely in the
thin clay layer located between the yellowish clay and the phos-
phatic clayey sand.
20 S. Chakir et al. / Cretaceous Research 106 (2020) 104219
Three main marine paleoenvironments are described in seven
intervals in this paper. An in inner neritic to rarely middle neritic
environment alternated with several short phases of more offshore
conditions e possibly related to the oceanic paleocurrents or a
decrease in nutrients due to a decrease of upwelling strength or
freshwater input e is recognized for the deposits of the upper
Maastrichtian and of the first ~130 cm above the K/Pg boundary.
The lowermost Danian deposits records the most pronounced
relative sea level fall in the section. And an open outer neritic
marine environment during a transgressive regime is determined
for the uppermost Danian part of the section.
We identified, using previous palynological studies of several
sections in the Tethyan realm, a succession of seven bioevents
inferred from ecological responses of the dinoflagellate cysts to
possible major climatic changes. These events include an episode of
warm climate conditions inferred from high abundances of the
tropical thermophilic Palaeocystodinium group (Andalusiella spp.
and Palaeocystodinium spp.) in the upper Maastrichtian, suspected
to be associated to a Late Cretaceous global warming event, prob-
ably related to Deccan traps volcanism. The warming event is fol-
lowed by an episode of relatively cold-water conditions deduced
from the incursion of the high-latitude cold-water Isabelidinium
group (mainly Isabelidinium cooksoniae), possibly related to the
latest Maastrichtian cooling that reflects the aftermath of the Late
Cretaceous global warming related to the Deccan volcanism. The
lowermost Danian sediments record cold-water conditions, indi-
cated by an abundance spike of the Isabelidinium group, which may
coincide with the episode of the first cooling pulse, related to the
well-known brief “impact winter” characterizing the first months
to decades following the Chicxulub impact. This episode is directly
followed by high-productivity conditions, denoted by a strong in-
crease in abundance of the Senegalinium group (mainly Sene-
galinium spp.). And then two other successive acmes of the
Palaeocystodinium group and Isabelidinium group, which may be
related respectively to the global warming following the K/Pg
boundary as a result of the release of greenhouse gases into the
atmosphere, followed by a second cooling pulse following the
initial cooling pulse at the K/Pg boundary. The uppermost part of
the section records a return to stable marine conditions, inferred
from a dominance of the gonyaulacoid groups, including the
cosmopolitan Spiniferites group (mainly Spiniferites spp.).
Acknowledgments
The authors thank the staffs of the Laboratory of Dynamics of
Sedimentary Basins and Geological Correlations (Faculty of Sci-
ences, Ben M'sik, Casablanca) and the Laboratory of Geo-
Biodiversity and Natural Heritage (“Geophysics, Natural Patri-
mony and Green Chemistry” Research Center, Scientific Institute,
Rabat) for their technical help. The first author S. Chakir thanks
“Centre National de la Recherche Scientique et Technique” (CNRST)
in Morocco for financial support during this study as a part of her
PhD. Thesis. This study is led by H. Slimani. The journal anonymous
reviewer, Eduardo Koutsoukos (Editor-in-Chief) and John W.M. Jagt
(Associate Editor) are kindly thanked for their critical and
constructive reviews that improved considerably the initial
manuscript. Their remarks are much appreciated.
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Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.org/10.
1016/j.cretres.2019.104219.