Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238

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Palaeogeography, Palaeoclimatology, Palaeoecology

journal homepage: www.elsevier.com/locate/palaeo

Marine vertebrate faunas from the Maastrichtian phosphates

of Benguérir (Ganntour Basin, Morocco): Biostratigraphy,
palaeobiogeography and palaeoecology
Henri Cappetta a,⁎, Nathalie Bardet b, Xabier Pereda Suberbiola c, Sylvain Adnet a, Driss Akkrim d,
Mohamed Amalik d, Aziza Benabdallah e
a
UMR 5554 du CNRS, Institut des Sciences de l'Evolution, Université Montpellier II Sciences et Techniques du Languedoc, Case courrier 064, place Eugène Bataillon, 34095 Montpellier cedex 5, France
b
Sorbonne Universités, CR2P, CNRS-MNHN-UPMC, Département Histoire de la Terre, Muséum National d'Histoire Naturelle, CP 38, 8 rue Buffon, 75005 Paris, France
c
Universidad del País Vasco/Euskal Herriko Unibertsitatea, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y Paleontología, Apartado 644, 48080 Bilbao, Spain
d
Office Chérifien des Phosphates, Centre Minier de Benguérir, Benguérir, Morocco
e
Ministère de l'Energie, des Mines, de l'Eau et de l'Environnement, Direction de la Géologie, Rabat, Morocco

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

Article history: The Maastrichtian of Benguérir (eastern part of the Ganntour Basin, Morocco) consists of about 20 m of
Received 13 September 2013 phosphates displaying an alternance of soft phosphate levels, marly horizons and hard phosphatic limestones.
Received in revised form 15 April 2014 Isolated teeth of selachians, actinopterygians and marine reptiles are extremely numerous in these phosphatic
Accepted 30 April 2014
deposits and have been used for biostratigraphical, palaeodiversity and palaeoecological purposes.
Available online 9 May 2014
Detailed field work allowed to establish an exhaustive list of the Benguérir marine vertebrate faunas with their
Keywords:
biostratigraphical distribution through five main fossiliferous levels (L6 to L2) spanning all the Maastrichtian.
Selachians Their importance for biochronological purposes and correlations with other Maastrichtian phosphate deposits
Actinopterygians worldwide appears noteworthy.
Reptiles The selachians are currently represented by 60 species belonging to 32 genera and 7 orders. Among them, the
Maastrichtian genus Squalicorax is one of the most interesting concerning high-resolution biostratigraphy and correlations
Morocco with other phosphate basins because of important rates of change noted between the 5 species recovered
Biodiversity from base (e.g. occurrence of S. africanus) to top (e.g. strong representation of S. pristodontus) of the
Maastrichtian. The marine reptiles include mainly mosasaurids but also scarcer plesiosaurs, chelonians and
crocodyliforms, representing at least 14 taxa. The mosasaurid squamates are the most abundant and diversified
with at least 8 species ranging all along the succession. The actinopterygians include mainly teleosts but also
pycnodonts, also common in all levels and representing at least 7 taxa.
Selachians and reptiles show the same trends, in terms of species richness per level, even if the reptiles are less
informative due to a less diversified assemblage. For sharks, L6 and L2 show a high percentage of genera and
species occurring only in the layer concerned. The evolution of diversity in actinopterygian fishes is less clear be-
cause of their low diversity. The use of dissimilarity indices and agglomerative method underscores two distinct
associations: a lower one including the levels L6 and L5, and an upper one comprising the levels L4 to L2. These
two associations allow to separate a lower and an upper Maastrichtian level and are important for correlations all
around the southern and eastern margins of the Tethys. Another clear faunal turnover occurs between L3 and L2,
because of a high appearance rate in L2 (at least in sharks) suggesting an increase in prey abundance, as testified
by the rapid increase of marine predator density.
Indeed, and through L6 to L2, a possible signal of an environmental damage affecting the predator community can
be noted by faunal turnovers, even if no significant change in prey association was clearly detected.
From a palaeobiogeographical point of view, the faunal associations of Benguérir appear typical of the southern
and eastern margins of the Tethys, with several typical species not occurring in the northern Tethys.
© 2014 Published by Elsevier B.V.

1. Introduction

⁎ Corresponding author.
E-mail addresses: henri.cappetta@univ-montp2.fr (H. Cappetta), bardet@mnhn.fr
(N. Bardet), xabier.pereda@ehu.es (X. Pereda Suberbiola), sylvain.adnet@montp2.fr
(S. Adnet), d.akkrim@ocpgroup.ma (D. Akkrim), m.amalik@ocpgroup.com (M. Amalik).
In the framework of an active French–Morrocan scientific collabora-
tion, many palaeontological field campaigns have been made during the
last decades in the Upper Cretaceous-Paleogene phosphatic basins of

http://dx.doi.org/10.1016/j.palaeo.2014.04.020
0031-0182/© 2014 Published by Elsevier B.V.
218 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238
Morocco, mainly in the Ouled Abdoun and Ganntour ones. Abundant
fossil vertebrate remains were collected here, including selachians
(Cappetta, 1981, 1983, 1984, 1985, 1986a, 1986b, 1986c, 1987, 1988,
1989, 1992; Noubhani and Cappetta, 1992, 1995, 1997, 2002),
actinopterygians (Cavin et al., 2000), reptiles (including birds) (Bardet
et al., 2010), and mammals (Gheerbrant et al., 2003). These new discov-
eries significantly improved our knowledge of these vertebrate assem-
blages, previously either unknown or represented by fragmentary or
isolated remains only (Arambourg, 1952).
Selachian, actinopterygian and mosasaurid remains are the most
abundant and diversified vertebrate fossils in the phosphatic series of
Morocco. Of particular importance are isolated teeth of these taxa,
which are well preserved and most often diagnostic at the generic and
even specific levels.
On the basis of a large and representative sample of isolated teeth
(several thousands), the aim of the present work is to provide the first
complete elasmobranch, bony fish and marine reptile faunal list of the
Benguérir series (Ganntour Basin), with their biostratigraphical distri-
bution, and to discuss their relevance for:
– biochronological purposes, especially relevant in the phosphatic
deposits where classical biomarkers (cephalopods, calcareous
nannofossils, foraminifera) are usually lacking, and correlations
with other phosphatic deposits in the world;
– faunal turnovers;
– and palaeobiogeographical distributions.
2. Abbreviations
BEG is the acronym for the Benguérir locality. The stratigraphical
level of each specimen is included directly in the collection number
as “C6” (Level 6) to “C2” (Level 2). MNHN, Muséum National d'Histoire
Naturelle (Paris, France); OCP, Office Chérifien des Phosphates
(Khouribga, Morocco).
3. Geographical and stratigraphical settings
During the Late Cretaceous and Early Paleogene, platforms border-
ing the Afro-Arabian Shield (southern Tethys and South Atlantic)
were characterised by phosphatic sedimentation in warm and
shallow-marine environments (Lucas and Prevot-Lucas, 1996). Phos-
phatic deposits, forming part of the so-called Mediterranean Tethyan
phosphogenic Province, currently crop out from North Africa to the
Near East where they constitute an economically valuable resource.
The phosphatic deposits of Morocco, known since 1908, have been
exploited since the 1920s (Office Chérifien des Phosphates, 1989).
They crop out in several basins, the main ones being the Ouled Abdoun
and the Ganntour basins, located between Casablanca and Marrakech
(Fig. 1A). Stratigraphically, these phosphatic strata range from upper-
most Cretaceous (lowermost Maastrichtian) to basal mid-Eocene
(Lutetian), spanning the longest time interval of all Tethyan phosphate
deposits (Lucas and Prevot-Lucas, 1996).
From a palaeogeographical point of view, the phosphates of the
Ouled Abdoun and Ganntour basins were deposited in a long and nar-
row gulf which opened west to the Atlantic Ocean (Herbig, 1986;
Salvan, 1986). The condensed succession in the northeastern part of
the Ouled Abdoun Basin probably was deposited in a high-energy, near-
shore environment located close to the Paleozoic Moroccan Massif
Central. The more complete sequences in the southwestern Ouled
Abdoun Basin and in the Ganntour Basin may have been laid down in
a more subsident, marine setting (Office Chérifien des Phosphates,
1989; Lucas and Prevot-Lucas, 1996).
The phosphate ore of Benguérir is located on the Ganntour Plateau,
about 12 km east of the Benguérir locality, north of the El Kelaa des
Sraghna road (Fig. 1B). It is limited by the Paleozoic Rehamna range
northwards and by the Paleozoic Djebilet range southwards.
The phosphatic series is intensively exploited in long north–south
oriented trenches. The Benguérir succession consists of about 20 m of
phosphate deposits displaying an alternance of soft phosphate levels,
marly horizons and hard phosphatic limestones. Contrary to the Ouled
Abdoun Basin where the phosphatic series spans the Maastrichtian
and Paleogene, most of the exploited phosphatic series in the Ganntour
Basin and especially in the Benguérir area is Maastrichtian in age.
4. Description of the Benguérir section
The Maastrichtian deposits can be observed in large and deep man-
made trenches oriented north–south, on a length of more than ten
kilometres, from the northern margin of the Benguérir plateau to the
road going to El Kelaa des Sraghna. Since few years, mining operations
have started south of this road.
Several soft phosphatic levels rich in vertebrate remains occur in the
succession, and five of them, called Levels 6 to 2 (L6 to L2, corresponding
to “couche 6” through “couche 2” in French) from the base to the top,
were particularly sampled (Fig. 2). These levels are briefly described
below. The total thickness of the phosphatic succession between the
base of Level 6 and the top of Level 2 is about 19 m. The thickness of
the “Sillon X” (not figured) is difficult to measure precisely because
the conditions of the outcropping at the top of the trenches, but can
be estimated at about 1.5 m. The “Sillon X” corresponds to the upper-
most Maastrichtian level of the Benguérir phosphatic series in the no-
menclature of the mine-workers.
The geology and stratigraphy of the Ganntour basin were mainly
studied by Boujo in a series of papers (1968, 1976, 1980). Below the
phosphate member starting with Level 6, there is a series of white and
yellow clays and sandstones, about 45 m thick and devoid of fossils.
This pre-phosphatic series, only known by boreholes, is still considered
as Maastrichtian in age (Boujo, 1976). However, it is not impossible that
the lower part of the pre-phosphatic series represents the Upper Cam-
panian. Anyway, the base of the phosphate series can be considered of
Early Maastrichtian age, following the conclusions of Boujo (1976). So,
it is highly likely that Level 6 represent the Lower, but not the lower-
most Maastrichtian. This assumption is supported by recent studies on
oxygen isotope measurements of shark teeth biogenic apatites from
the different levels of Benguérir (Kocsis et al., 2014).
The phosphatic succession overlies thick yellow marls and clays de-
void of fossils. It starts with level L6. The contact with the underlying
yellow marls is marked by a strongly bioturbated surface where one
can see, in places, an accumulation of large teeth, mainly Squalicorax
spp. and Scapanorhynchus rapax (Quaas, 1902). The phosphate at this
level is between 30 and 40 cm thick, coarse, very rich in fossil remains,
mainly actinopterygian fishes, with pebbles of phosphatic limestone,
marl and siliceous marl. Coarse siliceous sand is also particularly abun-
dant, underlining the beginning of the Maastrichtian transgression.
Above, there is a bed of marl and siliceous marl and chert, of 1.14 m
thick.
An intercalary L6/L5 level 66 cm thick consists of siliceous limestone
and marl, chert and coarse phosphate at the base.
Level L5 is 2.60 m thick. It exhibits an alternance of sandy phosphate,
chert and phosphatic limestone. The top of L5, about 1.90 m thick, con-
sists of marl, siliceous marl, sandy phosphate, with siliceous limestone
in places. Compared with the other sampled levels, L5 was particularly
poor in Squalicorax teeth, unlike to the other levels. For this reason, it
was not meaningful to take it into consideration for the analysis of the
Squalicorax abundance through the succession (Fig. 5).
The intercalary L5/L4 level is about 80 cm thick. It shows marly phos-
phate, phosphatic sand and phosphatic limestone, with siliceous marl in
places.
The L4 level is 5.12 m thick with about 2 m of marl, siliceous marl,
siliceous limestone and chert at its base. The phosphate bed is about
1.65 m thick. It consists of coarse sandy phosphate at its base with
chert and limestone; the phosphate is finer toward the top. Above,
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238
A 8°
Casablanca
Atlantic
Ocean
Safi Youssoufia
32° Benguérir
GANNTOUR
Marrakech
B Casablanca
“Tranchée d’essai”
Benguérir (L6)
Marrakech 0 10 km
Fig. 1. Location map of the studied sites in Morocco. A: Ganntour and Ouled Abdoun Basins. B: Sampled area (grey rectangle) east of Benguérir, central part of the Ganntour Basin.
there is a level 1.42 m thick consisting mainly of marl and phosphatic
limestone.
Level L3 is 1.50 m thick. It consists of a basal level of sandy phosphate
and an upper level of phosphatic sand.
Above, one can observe a very typical level of yellow clay about 1.90
m thick which is very constant in the basin. The top is bioturbated and
the burrows are infilled by phosphate from the level L2.
Level L2 shows two phosphate levels. The lower one is about 1.30 m
thick; it consists of phosphatic sand with, in places, phosphatic lime-
stone. The upper one, about 1.40 m thick, is a coarse phosphate rich in
organic remains. The two phosphate levels are separated by about
50 cm of phosphatic limestone, marl and clay.
The “Sillon X” (SX) could not be identified with certainty above the
L2 level. Several thin and rather lenticular phosphate levels occur at
the top of the succession, mainly in the south of the mining area. Yet,
their faunal content seems to be a mixture of Maastrichtian and Danian
elements, as that can be observed in the Ouled Abdoun and contrary to
the western part of the Youssoufia area where the SX can be separated
by its faunal content (Cappetta, 1987). So, further investigations are
necessary to try to solve this problem in the Benguérir area.
219
.
Khouribga
Oued Zem
OULED
ABDOUN
0 50 km
Beni Mellal
El Kelaa des
Sraghna
Marrakech
5. Material and methods
The first vertebrate fossils were obtained by the senior author in
1979, just before the start of industrial mining in the 1980s. At this
time, only some parts of the phosphatic series were accessible in small
open pits, and only the L6 could be properly sampled, just north of the
road Benguérir–Marrakech (Noubhani and Cappetta, 1997 and Fig. 1B,
“Tranchée d'essai” site). Despite the richness and diversity of the verte-
brate fauna, consisting mainly of elasmobranch teeth but also including
abundant marine reptile remains, no study has been dedicated to this
locality until now. Noubhani and Cappetta (1997) described the faunal
content of L6, that was available at the time, on the basis of the samplings
of the senior author. The field-trips made since 1998 have allowed to
complete the sampling of all the vertebrate-bearing horizons.
The fossils were obtained partly by surface collecting in the trenches,
but also on areas where the different phosphate levels are kept in stock
separately and not mixed. During the mining process the different phos-
phate levels are removed in successive bearings, from the top down to
the lowest Level 6, in broad and very long man-made trenches. There-
fore, it has been possible to collect teeth in the successive levels, on
220 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238

LEVELS LITHOLOGY
(Phosphate)

1.40 m
L2
upper
0.52 m

1.30 m L2
lower

1.86 m
yellow clays
(marker level)

1.50 m L3

1.42 m

1.65 m L4
2s

2.05 m
Yellow clay

. . . . . . . . . .
intercalary
. . . . . . .
. . . . . . . . .
. . . . . . . . .
Marls and siliceous limestone
0.80 m . . . . . . . ... . .
. . . . .. . . . . .
L5/L4 . . . . . . . ..
. . . . . . .
. . . . . . . .. . . .. . .
.
.
. .
. . . .
. . . . .
. . . . .
Sandy phosphate,
.. . .
. .
.
. . . . .
. + ++++
. . . . .
limestone and silex
1.90 m . . . . .. . . . .
. . . .. . .
Calcareous phosphate

Marls

2.60 m L5
Soft phosphate

Yellow marls
0.66 m intercalary +++

L6/L5 +
+ + ++
+ ++ + + +
Soft phosphate and
+
calcareous phosphate
++ + + + +
1.14 m L6

0.35 m

Fig. 2. Simplified composite section of the Maastrichtian phosphatic series East of Benguérir.
Modified from L. Bensalah and M. Aguenagay.
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238
very wide areas corresponding each time to a single level. Furthermore,
the phosphate extracted from a precise level is placed separately in stor-
age areas, without being mixed with phosphate coming from other
levels, avoiding thus mix-up or contamination. This particularity allowed
us to collect numerous teeth on these stocks from identified levels,
supplementing the samplings made in the trenches. Large amounts of
sediment from the different levels were also sieved, using mesh sizes
down to 400 μm, in order to collect the micro-ichthyofauna. Sieving
has not been performed for mosasaurids as the juvenile teeth are rela-
tively uninformative.
Due to their abundance and taxonomical utility, attention has been
paid mainly to selachian, actinopterygian and mosasaurid teeth. In each
level of the Maastrichtian phosphatic succession, the number of taxa
has been taken into account, permitting diversity comparisons between
each phosphatic levels. For each taxon, a stratigraphical range has been
proposed. Lazarus taxa have been taken into account when elaborating
diversity analyses and curves. For some groups (mosasaurids, selachians
especially Squalicorax) the number of specimens (abundance) has also
been considered. For palaeoecological and palaeobiogeographical pur-
poses, comparisons have been made with available data obtained from
the nearby located Ouled Abdoun Basin but also from other phosphatic
deposits worldwide, especially those from the Middle East. All these
data, especially those based on the selachian assemblages at each level,
have improved our understanding of, not only the age of the Morrocan
phosphates but also that of the other phosphatic deposits from the
southern margin of the Tethys.
For palaeoecological purposes, agglomerative analysis on dissimilar-
ity indices (Raup and Crick, 1979) has been performed.
6. Vertebrate diversity and stratigraphical range
6.1. Selachians
Their remains, essentially isolated teeth, are by far the most abun-
dant vertebrate fossils at all levels of the phosphatic series. A total of
60 species belonging to 32 genera have been identified. Table 1 summa-
rizes the faunal content of the different levels. Fig. 3 shows the generic
(Fig. 3A) and specific (Fig. 3B) total diversity. Fig. 4 shows details
order by order of the generic (Fig. 4A) and specific (Fig. 4B) diversity. Fi-
nally, Figs. 6 and 7 provide illustrations of some key taxa.
The diversity and abundance of the different assemblages vary
through all the succession, from L6 to L2. As shown in Fig. 3, the generic
diversity rises and falls in tandem with species diversity. In L6, the elas-
mobranch association is diversified both at genus and species levels.
There is a marked drop in diversity between L6 and L5, and after, an in-
crease in this diversity, mainly between L3 and L2.
Hexanchiformes and Pristiophoriformes only occur in the L2 assem-
blage (Fig. 4A–B, Table 1). The Hexanchiformes are represented in L2 by
scarce and fragmentary teeth of Hexanchus microdon (Agassiz, 1835).
The same applies to the Pristiophoriformes, which are represented by
very few specimens of rostral and oral teeth of Pristiophorus sp. Com-
bined, these remains could be indicative of deeper conditions of the
environment.
The Lamniformes are, by far, the most abundant and diversified
group among sharks with 7 genera and up to 18 species (Fig. 4A–B,
Table 1). However, this apparent abundance is partly a bias due to the
large size of the teeth that can be collected in large numbers by surface
picking. It is more difficult to collect small to very small teeth of other
groups, like the Rajiformes and many Myliobatiformes, obtained by
fine mesh sieving (down to 0.4 mm). However, this collecting bias
does not influence the number of taxa in each level. Lamniformes in-
clude a maximum of six genera and nine species in L2. L6 is even richer,
with 10 species in four genera. The higher diversity in L6 is mainly due
to the occurrence of species exclusive to this level, as for example
Scapanorhynchus rapax and Cretolamna biauriculata. At the specific
level, this order is as diversified as is the Myliobatiformes in the whole
221
succession, except in L5 where Lamniformes are twice as diverse. But
this situation is probably a bias due the relative poverty of this level.
Throughout the stratigraphic sequence, Lamniformes and in particu-
lar Squalicorax and Cretolamna are especially abundant. Only some
Lamniformes taxa, particularly in Squalicorax, have been considered in
terms of relative abundance through the stratigraphic series.
Interesting variations of diversity can be observed in the genus
Squalicorax (Pomes, 2003). The genus is abundant in all levels of the suc-
cession, except in L5, but the stratigraphic range and abundance of the
different species greatly vary (Fig. 5). The L6 level contains the most di-
verse specific assemblage of this genus, with five species, three of them
representing a large portion of the specimens: S. pristodontus (43%), S.
nov. sp. 1 (38%) and S. nov. sp. 2 (17%). Squalicorax bassanii constitutes
only 2% of the assemblage. Two species, S. africanus and S. nov. sp. 2, do
not occur above L5. In unit L4, S. pristodontus dominates the assemblage
with more than 90% of the teeth and almost all anacoracid teeth in L2
(99%) can be assigned to this species. The species S. africanus is not rep-
resented in Fig. 5 because of the very small size of its teeth. Indeed, be-
cause the Squalicorax species teeth were obtained mainly by surface
collecting, the number of teeth of S. africanus is underrepresented com-
pared to the other species with large sized teeth. The species S. bassanii
and S. nov. sp. 1 are present from L4 to L2 but represent a low percent-
age of the Squalicorax association (Fig. 4). Squalicorax africanus (Fig. 7D–
E), a very small-sized teeth species, is particularly important for correla-
tions with other southern Tethyan phosphate basins. Squalicorax
bassanii (Fig. 7A–C), a species misidentified by Arambourg (1952) as
S. kaupi, is present through all the succession but its abundance greatly
varies according to the levels. It is not scarce in L6, and seems to be par-
ticularly common at the base of this level. It is present from L4 to L2, but
becomes scarcer upsection, coupled with a clear size increase.
Squalicorax nov. sp. 2 (a species described in a separate paper in prepa-
ration and corresponding to S. yangaensis in Arambourg, 1952) is com-
mon in L6; one fragmentary tooth comes from L5 but the species
disappears completely above this level. This species is important for cor-
relations with the Middle East. Squalicorax nov. sp. 1 is also common in
L6. It can be collected up to L3 where it becomes scarce. Squalicorax
pristodontus (Fig. 7F–I) is the only species occurring at all levels of the
succession. It is rather abundant everywhere but seems to be scarcer
in L2 even if it comprises 99% of the Squalicorax association in this
level. It is interesting to note that specimens of this lineage from L6
are actually much closer to S. lindstromi (Davis, 1890) than they are
close to the S. pristodontus population in L2 (M. Siversson, in litt.). More-
over, one can note some morphological changes of the teeth between L6
and L2. At the base of the succession, the teeth are more asymmetrical,
with a clear notch separating the distal heel from the distal cutting edge
of the cusp. In L2, the teeth are much more symmetrical. These morpho-
logical changes are also important for correlations. It is likely that a de-
tailed study of the successive populations of S. pristodontus through the
Benguérir succession will lead to the splitting of this species into several
chrono-species.
The genus Pseudocorax is only represented in L2 by a single tooth of
P. affinis. This genus is, without exception, very scarce in the Ganntour
and Ouled Abdoun basins. It is only rather common in the Maastrichtian
of the Bekrit area (Middle Atlas, H.C. unpubl. data). This taxon is proba-
bly indicative of a more open sea or deeper environment.
Scapanorhynchus rapax (Fig. 6A–D) is a noteworthy element of the
Benguérir fauna due to the large size of its teeth (up to 6 cm high for an-
terior teeth). It was one of the largest predators among sharks of the
Maastrichtian phosphatic sea, besides Cretolamna maroccana and
Squalicorax pristodontus. It is particularly abundant just at the base of
L6 and becomes scarcer higher up within L6. It is absent above L6.
Anomotodon plicatus teeth are not very common, as is the case in the
Ouled Abdoun Basin, and occur only at the top of the succession, in L2.
The genus Cretolamna, with five different species, is one of the most
abundant genera in terms of tooth numbers. The species C. maroccana
(Fig. 6I–K) occurs from L4 to L2 but is absent in L6 and L5 where it is
222 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238

Table 1
List of selachians, reptiles and actinopterygians taxa identified in the Maastrichtian Benguérir succession, from L6 to L2. SHA: sharks; BAT: batoids; REP: marine reptiles; BFI: bony fishes.
See Fig. 15B for explanation of numbers corresponding to cutting and grinding abilities.

Ref Subgroups Species Level 6 Level 5 Level 4 Level 3 Level 2 Tooth size Cutting ability Grinding ability
(cm)

1 SHA Hexanchus microdon X 1 2 1

2 SHA Pristiophorus sp. X 0.4 0 1
3 SHA Squalicorax africanus X X 0.6 3 0
4 SHA Squalicorax bassanii X X X X X 3 3 0
5 SHA Squalicorax nov. sp. 1 X X X X 2 3 0
6 SHA Squalicorax nov. sp. 2 X X 2 3 0
7 SHA Squalicorax pristodontus X X X X X 4 3 0
8 SHA Pseudocorax affinis X 1 2 0
9 SHA Scapanorhynchus rapax X 6 1 0
10 SHA Anomotodon plicatus X 2 1 0
11 SHA Cretolamna appendiculata X 2 1 0
12 SHA Cretolamna maroccana X X X 3 2 0
13 SHA Cretolamna aff. maroccana X X 2 2 0
14 SHA Cretolamna biauriculata X 1.5 2 0
15 SHA Cretolamna sp. X 1.5 2 0
16 SHA Serratolamna africana X 3 1 0
17 SHA Serratolamna serrata X X X 4 1 0
18 SHA Serratolamna khderii X X 2 1 0
19 SHA Serratolamna aff. khderii X X 2 1 0
20 SHA “Carcharias” heathi X 2 1 0
21 SHA Ginglymostoma sp. X 1 1 0
22 SHA Ganntouria variabilis X 1 1 0
23 SHA Plicatoscyllium lehneri X 0.5 1 0
24 SHA Plicatoscyllium youssoufiaense X 0.5 1 0
25 SHA Plicatoscyllium gharbii X X X X 0.5 1 0
26 SHA Plicatoscyllium pectinatum X 0.5 1 0
27 SHA Plicatoscyllium sp. X 0.5 1 0
28 SHA Chiloscyllium sp. 1 X 0.4 1 1
29 SHA Chiloscyllium sp. 2 X 0.3 1 1
30 SHA Scyliorhinus aff. elongatus X 0.3 1 0
31 SHA Pteroscyllium sp. X X 0.4 1 0
32 BAT “Rhinobatos” sp. 1 X X X 0.2 1 1
33 BAT “Rhinobatos” aff. sp. 1 X 0.2 1 1
34 BAT “Rhinobatos” sp. 2 X X 0.2 1 1
35 BAT “Rhinobatos” sp. 3 X 0.2 1 1
36 BAT “Rhinobatos” sp. 4 X X 0.2 1 1
37 BAT “Rhinobatos” sp. 5 X 0.2 1 1
38 BAT “Rhinobatos” aff. primarmatus X 0.3 1 1
39 BAT Hypsobatis weileri X X 0.2 0 2
40 BAT Hypsobatis sp. X 0.5 0 2
41 BAT Youssoubatis aff. ganntourensis X X 0.5 0 2
42 BAT Youssoubatis sp. X 0.4 0 2
43 BAT Parapaleobates atlanticus X 0.2 0 3
44 BAT Rhinobatoidei indet. X 0.3 0 2
45 BAT Dalpiazia stromeri X X X X X 0.4 0 0
46 BAT Ganopristis leptodon X X X X X 0.3 0 0
47 BAT Ctenopristis nougareti X X X X X 0.2 0 0
48 BAT Schizorhiza stromeri X 0.3 0 0
49 BAT Duwibatis sp. X 0.3 0 0
50 BAT Dasyatis sp. 1 X 0.2 1 1
51 BAT Coupatezia elevata X X X 0.2 1 2
52 BAT Coupatezia fallax X 0.2 1 2
53 BAT Coupatezia sp. X 0.2 1 2
54 BAT Dasyrhombodus bondoni X X 0.3 1 3
55 BAT Dasyatoidea nov. gen. nov. sp. X X 0.2 0 3
56 BAT Ixobatis mucronata X X 0.3 0 3
57 BAT Rhombodus binkhorsti X 0.4 0 3
58 BAT Rhombodus microdon X X X X 0.3 0 3
59 BAT Rhombodus meridionalis X X X X 0.3 0 3
60 BAT Cretomanta sp. X X X X 0.2 1 2
61 REP Pachyvaranus crassispondylus X X X X 1 0 0
62 REP Mosasaurus beaugei X 6 2 1
63 REP “Platecarpus” ptychodon X X X X X 2.5 1 0
64 REP Halisaurus arambourgi X X X X X 3 0 0
65 REP Globidens phosphaticus X X X X 4 0 3
66 REP Carinodens belgicus X 3.5 0 2
67 REP Prognathodon giganteus X X 6.5 2 1
68 REP Prognathodon sp. X X X 6 2 1
69 REP Prognathodon currii X X X X X 6 2 2
70 REP Eremiasaurus heterodontus X X X X X 6 2 2
71 REP Elasmosauridae indet. X X X X 4 0 0
72 REP Bothremydidae indet. X X X X X
73 REP Chelonioidea indet. X X X X X
74 REP Dyrosauridae indet. X X X 1.5 1 0
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238 223

Table 1 (continued)
Ref Subgroups Species Level 6 Level 5 Level 4 Level 3 Level 2 Tooth size Cutting ability Grinding ability
(cm)

75 REP Eusuchia indet X 1.5 1 0

76 BFI Enchodus elegans X X X 3 1 0
77 BFI Enchodus libycus X X X 6 1 0
78 BFI Enchodus bursauxi X X X 4.2 1 0
79 BFI Stratodus apicalis X X X X X 0.5 1 0
80 BFI Stephanodus libycus X X 0.6 1 1
81 BFI Pseudoegertonia sp. X X 0.4 0 3
82 BFI Pycnodontidae indet. X X 0.8 0 3

replaced by a dentally similar species reminding C. maroccana, probably
representing a new species (here referred as C. aff. maroccana).
Cretolamna biauriculata (Fig. 7J–L) is restricted to L6, as is Cretolamna
sp. (Fig. 6L–O), which shows a very distinct tooth morphology (thick
root; thick, slightly twisted crown; broad lateral cusplets, often divided
into two), indicating that it is a new species. The occurrence of
C. appendiculata rests on some teeth collected in L2. These teeth are
very similar to specimens collected higher in the series, in the Danian,
and it is likely that they represent an ancestor of the Danian species.
The genus Serratolamna is well represented with three species
showing different stratigraphical ranges. Serratolamna africana
(Fig. 6G–H), an uncommon small-sized species, occurs only in L2.
Serratolamna serrata (Fig. 6E–F) occurs from L4 to L2 and the tooth
size increases between these two levels. Serratolamna khderii is a species
with anterior teeth of odontaspidid morphology and with lateral teeth
morphologically close to those of the species S. serrata, but with cusps
bearing folds on the lingual face. The species S. khderii has been defined
from the Early Maastrichtian of Jordan (Zalmout and Mustafa, 2001)
and it occurs in Benguérir in L6 and L5. Teeth resembling those of
S. khderii, but larger in size, occur higher in the succession (L4 and L3);
it is possible that some of those teeth belong to a new species, here pro-
visionally referred to as S. aff. khderii.
“Carcharias” heathi is an odontaspidid that only occurs in L2; it is of
rather large size, with a slightly heterodont dentition, compared to the
usually clearly heterodont dentition of odontaspidids. The taxon also oc-
curs in Level III of the Ouled Abdoun Basin, which is equivalent to L2 in
the Ganntour Basin.
The Orectolobiformes are represented by four genera. The maximum
diversity is observed in L2 with four species. An indeterminate
Ginglymostoma species with poorly preserved teeth occurs in L3. The
genus Plicatoscyllium is abundant, mainly in L6, with P. gharbii, which
occurs up to L3. Two other species, P. youssoufiaense and P. aff.
pectinatum are present only in L2. Plicatoscyllium lehneri is also repre-
sented in L2 by several incomplete but typical teeth. Moreover, teeth
of Ganntouria variabilis occur only in L2. Two uncommon species of
Chiloscyllium with very small teeth have been collected in L6.
The Carcharhiniformes, little diversified and very scarce, are only
represented by the scyliorhinid genera Pteroscyllium sp., in L6 and L2,
and Scyliorhinus aff. elongatus in L2.
The batoid rays (Rajiformes, Myliobatiformes) are abundant at all
levels of the Benguérir succession, with at least 17 genera and 29 spe-
cies. The genera of Rajiformes are diversified from L6 and show the
same tendency as for Myliobatiformes, except for L2 where they are as
important as in L3. At the specific level, 11 species are present in L6,
four in L5, six in L4 and L3, and nine in L2. The genus “Rhinobatos” l. s.
is represented by seven different species. According to their dental mor-
phology, very different from the morphology of the examined recent
species of the genus, it is obvious that most of those fossil “Rhinobatos”
species correspond in fact to different genera, some of them probably
new. Therefore, the generic diversity of batoids is understated. Some
occur only at the base of the succession (L6): “R.” aff. primarmatus,
“Rhinobatos” sp. 5 and “Rhinobatos” aff. sp. 1. Some occur in the upper
part of the succession (L4 to L2): “Rhinobatos” sp. 1, “Rhinobatos” sp. 2
and “Rhinobatos” sp. 4. “Rhinobatos” sp. 3 is found only in L2.
Among the Hypsobatidae, Hypsobatis weileri is particularly abundant
in L6. A morphologically close, but yet different species is represented in
L3 by scarce specimens. Youssoubatis aff. ganntourensis occurs only at
the top of the succession, in L3 and L2. All the collected specimens of

A 25
Number of Taxa

20
Reptiles
15
Actinopterygians
10
Selachians
5
0
L6 L5 L4 L3 L2
Stratigraphical levels

B 35
Number of Taxa

30
25 Reptiles
20
Actinopterygians
15
10 Selachians
5
0
L6 L5 L4 L3 L2
Stratigraphical levels

Fig. 3. Total marine vertebrate diversity (selachians, actinopterygian fishes and reptiles) according to the different Maastrichtian phosphatic levels (L6 to L2) of Benguérir. A: generic di-
versity. B: specific diversity.
224 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238

A -Genera
100%

90%

80% Myliobatiformes
70% Rajiformes
60%
Carcharhiniformes
50%
Orectolobiformes
40%
Lamniformes
30%
Pristiophoriformes
20%
Hexanchiformes
10%

0%
L6 L5 L4 L3 L2

B -Species
100%

90%

80%
Myliobatiformes
70%
Rajiformes
60%
Carcharhiniformes
50%
Orectolobiformes
40%
Lamniformes
30%
Pristiophoriformes
20%
Hexanchiformes
10%

0%
L6 L5 L4 L3 L2

Fig. 4. Evolution of selachian ordinal diversity according to the different phosphatic levels (L6 to L2) of Benguérir, considering the specific and generic richness. A: Percentage of species per
order. B: Percentage of genera per order.

this last species are larger, with a higher crown, than are the type spec-
imens described from the “Sillon X” of Youssoufia (Cappetta, 1992).
Concerning the Parapalaeobatidae, Parapalaeobates atlanticus is
present only in L6 where it remains scarce.
The Sclerorhynchidae occur throughout the whole succession, ex-
cept for Schizorhiza stromeri which is collected only in L2. This family
is never abundant in the Ganntour succession, except Ctenopristis
nougareti, represented by hundreds of oral and rostral teeth mainly
from L6. Dalpiazia stromeri and Ganopristis leptodon are uncommon
but present through all the succession.
The genus Duwibatis, a genus defined in the Maastrichtian of Egypt
(Cappetta, 1991), is represented by some teeth in L6.
Seven genera and 11 species of Myliobatiformes have been
recognised in the Benguérir succession. They are well represented in
L6 with 4 genera, then they decrease to L5 (3 genera), increase again
in L4, L3 and L2 (4 genera). At the specific level, the tendency is similar,
except for L2 where they increase.
The genus Dasyatis is very scarce and represented by very few spec-
imens of small size in L2.
The genus Coupatezia is more abundant and represented by three
species: C. elevata in L4 to L2 and C. fallax in L2. A species with very
small-sized teeth is rather common in L6.
Some specimens of Dasyrhombodus bondoni (Arambourg, 1952)
have been identified in L4 and L3. This allows for a more precise
stratigraphical range determination. The species was originally de-
scribed from imprecise Maastrichtian levels of El Borouj (Ouled Abdoun
Basin) by Arambourg (1952).
An undescribed Dasyatoidea nov. gen. nov. sp. (under study) is well
represented at the base of the succession, in L6 and L5. This taxon is
widespread at the base of the Maastrichtian of many localities (Oued
Erguita and Bekrit in Morocco, Jordan, Syria and Egypt).
Ixobatis mucronata is observed only at the top of the succession, in L3
and L2.
The genus Rhombodus is represented through all the succession, but
unevenly. Three species are known, the largest one, R. binkhorsti
(Fig. 7N–Q) is abundant in L2 while R. meridionalis occurs in L6 and L5
(one tooth) and ranges up to L3 (one tooth); this last species is very
scarce in Benguérir, whereas it is very common at the base of
Maastrichtian of Oued Erguita (Souss Basin). R. microdon occurs in all
levels but L5.
The genus Cretomanta is represented by an indeterminate species
and occurs through all the Benguérir succession (L6- L3, but not in L2)
but remains always very scarce.
6.2. Actinopterygians
Bony fish remains are commonly found in the Maastrichtian
Benguérir succession, being represented mainly by isolated teeth,
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238 225

S. bassanii
1%
probably underestimated, due to the fact that a global review of these
bony fish has never been performed since the work of Arambourg
(1952).
The actinopterygians include mainly teleosteans but also
S. pristodontus
99% pycnodontids. All actinopterygians are present throughout the
Maastrichtian of Benguérir, at either the generic (Enchodus,
Stratodus, Stephanodus) or familial (phyllodontids, pycnodontids)
L2 systematical rank, except in “Sillon X”. These assemblages appear
thus very constant between the lower and the upper parts of the
Maastrichtian.
Enchodontids, represented by the genus Enchodus, are extremely
S. bassanii
1%
S. nov. sp. 1 abundant at all levels. They are present in the “Sillon X” but are here
5%
rare and of very small size. The three species described by Arambourg
(1952) have been identified on the basis of isolated teeth. Enchodus
was a large Late Cretaceous open-sea predator (Goody, 1976) and the
three species reported here are important for biostratigraphical correla-
tions. Indeed, Enchodus elegans (Fig. 8A) is very abundant in the lower
S. pristodontus
94% part of the succession (L6 to L4) but is apparently absent in the upper
part (L3 and L2). This distribution matches the one observed in the
Lower Maastrichtian phosphates of Syria, where E. libycus and
E. bursauxi are lacking (Bardet et al., 2000). In contrast, E. bursauxi
L3 (Fig. 8B) and E. libycus (Fig. 8C) are absent in the lower part of the suc-
cession (L6 to L4) but abundant and represented by very large teeth
(some can reach 10 cm in height) in the upper part (L3 and L2). They
S. bassanii S. nov. sp. 1 are present (though rare) in “Sillon X” and have also been found in
1% 5%
abundance in Level III (Upper Maastrichtian) of the Ouled Abdoun
Basin, where E. elegans is totally absent. The three species have thus
been found together only in L4. These taxa thus appear to be useful
biostratigraphical tools to discriminate between the lower (with
E. elegans) and the upper (with E. libycus and E. bursauxi) parts of the
Maastrichtian Benguérir phosphatic succession.
S. pristodontus
94% Remains of the dercetid Stratodus apicalis (Fig. 8D), represented by
jaw remains and vertebrae, are common throughout the Maastrichtian
of Benguérir, except in “Sillon X”. This large open-sea predatory teleost
is also present in Level III (Upper Maastrichtian) of the Ouled Abdoun
Basin.
L4 The eotrigonodontid Stephanodus libycus (Fig. 8E) is very scarce. Its
teeth have been found only in L6 and L2, but this distribution indicates
S. bassanii
that it is was present all throughout the Maastrichtian succession. It is
2% also present in Level III (Upper Maastrichtian) of the Ouled Abdoun
Basin.
Phyllodontid crushing teeth (Fig. 8G) have been only found in L6
S. nov. sp. 1
38% and L2, indicating that this taxon also spans the Maastrichtian suc-
S. cession. These teeth look like those of Pseudoegertonia previously
pristodontus
43% described from the Maastrichtian of Morocco and Middle-East, and
the Palaeocene of Congo (references in Bardet et al., 2000). They
are also present in Level III (Upper Maastrichtian) of the Ouled
Abdoun Basin.
Finally, pycnodontid isolated globulous teeth (Fig. 8F) are very
L6 scarce and have been found only in L6 and L2, indicating that they
S. nov. sp. 2
17% were present throughout the Benguérir succession, at least at the famil-
ial level. They are also present in Level III (Upper Maastrichtian) of the
Fig. 5. Diversity and relative abundance of the Squalicorax species through the Ouled Abdoun Basin.
Maastrichtian phosphate levels of the Benguérir series. Squalicorax africanus and L5 are
not represented (see text).

6.3. Reptiles

cranial bones and vertebrae. At least seven taxa are known. Table 1 sum-
marizes the faunal content of the different levels. Fig. 3 shows the gener-
ic (Fig. 3A) and specific (Fig. 3B) diversity. Fig. 8 provides illustrations of
the taxa.
The diversity of the actinopterygians throughout the Benguérir phos-
phatic succession is the lowest of all the three groups studied and also the
most constant, varying very little from L6 to L2. As shown in Fig. 3, the ge-
neric diversity is roughly comparable to the species diversity with around
five taxa for each level. However, the real actinopterygian diversity is
Marine reptile remains are abundant and represented by isolated
teeth, vertebrae and dermal scutes attributed mainly to mosasaurid and
pachyvaranid squamates and elasmosaurid plesiosaurs. Chelonians and
especially crocodyliforms remain particularly scarce in the Maastrichtian
phosphatic succession.
As selachians and bony fish, most species are present in all the stud-
ied levels but L5 and “Sillon X”, suggesting that the marine reptile as-
semblages, at least the mosasaurid ones were constant between the
Lower and the Upper Maastrichtian.
226 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238

A B C

E H
F

K
J
I

L M O

Fig. 6. Lamniform sharks from Benguérir, Ganntour Basin, Morocco. The stratigraphical level of each specimen is included directly in the collection number as “C6” (Level 6) to “C2” (Level
2). A–D: Scapanorhynchus rapax, A–C, anterior tooth, UM-BEG-C6.13; A, labial view; B, profile; C, lingual view; D, lateral tooth, UM-BEG-C6.14, lingual view. E–F: Serratolamna serrata; E,
anterior tooth, UM-BEG-C2.2, lingual view; F, lateral tooth, UM-BEG-C2.8, lingual view. G–H: Serratolamna africana; G, anterior tooth, UM-BEG-C2.13, labial view; H, lateral tooth, UM-BEG-
C2.14, lingual view. I–K: Cretolamna maroccana; I, anterior tooth, UM-BEG-C2.19, lingual view; J, lower lateral tooth, UM-BEG-C2.20, lingual view; K, upper lateral tooth, UM-BEG-C2.21,
lingual view. L–O: Cretolamna sp.; L–M, anterolateral tooth, UM-BEG-C6.44; L, profile; M, lingual view; N, lateral tooth, UM-BEG-C6.45, lingual view; O, very lateral tooth, UM-BEG-C6.46,
labial view.

At least 15 taxa have been identified. Table 1 summarizes the fau-
nal content of the different levels. Fig. 3 shows the generic (Fig. 3A)
and specific (Fig. 3B) total diversity. Fig. 8 provides illustrations of
the taxa.
The generic and specific diversity of marine reptiles is higher than
those of actinopterygians but lower than those of selachians (Fig. 3).
The diversity is also roughly constant at both the generic and specific
levels (around 10 taxa for each level) and through time (from L6 to
L2). Highest diversity is recorded in L6 and L2 levels (12 taxa), followed
by L4 and L3 levels (11 taxa). The low diversity of L5 is probably due to a
collecting bias as this level is generally very poor in fossils despite our
collecting efforts in this layer.
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238 227

Fig. 7. Lamniform sharks and myliobatiform rays from Benguérir, Ganntour Basin, Morocco. The stratigraphical level of each specimen is included directly in the collection number as “C6”
(Level 6) to “C2” (Level 2). A–C: Squalicorax bassanii; A–B, anterior tooth, BEG-C6.41; A, labial view; B, lingual view; C, lateral tooth, BEG-C6.42, labial view. D–E: Squalicorax africanus; D,
anterolateral tooth, BEG-C6.36, labial view; E, lateral tooth, BEG-C6.37, lingual view. F–I: Squalicorax pristodontus; F–G, anterior tooth, BEG-C2.17; F, labial view; G, lingual view; H, lateral
tooth, BEG-C2.18, labial view; I, lateral tooth, BEG-C3.3, lingual view. J–L, Cretolamna biauriculata; J–K, anterior tooth, BEG-C6.38; J, labial view; K, lingual view; L, lateral tooth, BEG-C6.39,
lingual view; M–N, Squalicorax nov. sp. 1; M, lateral tooth, BEG-C6.35, lingual view; N, detail of the mesial cutting edge, labial view. O–P, Squalicorax nov. sp. 2; O, lateral tooth, BEG-C6.24,
lingual view; P, detail of the mesial cutting edge, lingual view. Q–T, Rhombodus binkhorsti; Q–R, anterior tooth, BEG-C2.15; Q, lingual view; R, profile; S–T, lateral tooth, BEG-C2.16; S, oc-
clusal view; T, lingual view.
228 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238

Fig. 8. Actinopterygian fishes and marine reptiles (mosasaurid and pachyvaranid squamates and plesiosaurs) from Benguérir, Ganntour Basin, Morocco. A: Enchodus elegans, OCP-SA 653
(Level 6), palatine tooth, posterior view. B: Enchodus bursauxi, OCP-SA 654 (Level 2), palatine tooth, lateral view. C: Enchodus libycus, OCP-SA 655 (Level 2), palatine tooth, lateral view. D:
Stratodus apicalis, OCP-SA 656 (Level 3), jaw (dentary or maxilla) fragment, dorsal view. E: Stephanodus libycus, OCP-SA 657 (Level 2), pharyngeal tooth, lateral view. F: indeterminated
Pycnodontidae, OCP-SA 658 (Level 2), tooth, occlusal view. G: Pseudoegertonia sp., OCP-SA 659 (Level 6), pharyngeal plate fragment. H: Mosasaurus beaugei, OCP-SA 660 (Level 2), marginal
tooth, labial view. I: “Platecarpus” ptychodon, OCP-SA 661 (Level 3), marginal tooth, labial view. J: Halisaurus arambourgi, OCP-SA 662 (Level 2), marginal tooth, labial view. K: Eremiasaurus
heterodontus, OCP-SA 663 (Level 2), marginal tooth, labial view. L: Carinodens belgicus, G. Barbe's private collection (cast MNHN 6340, Grand Daoui, Ouled Abdoun Basin, Level CIII, Upper
Maastrichtian; see Bardet et al., 2008), marginal tooth, labial view. M: Globidens phosphaticus, OCP DEK-GE 361 (Holotype, Level 3; see Bardet et al., 2005b), marginal tooth, labial view. N:
Prognathodon currii, OCP DEK-GE 349 (Level 5), marginal tooth, labial view. O: Prognathodon giganteus, OCP-SA 664 (Level 6), marginal tooth, labial view. P: Prognathodon sp., OCP-SA 665
(Level 2), marginal tooth, labial view. Q: Pachyvaranus crassispondylus, MNHN PMC 21 (Level 4; see de Buffrenil et al., 2008), dorsal vertebra, ventral view. R: indeterminated
Elasmosauridae, tooth, OCP-SA 666 (Level 2), labial view.

The pachyvaranid squamate Pachyvaranus crassispondylus (Fig. 8Q) abundant in the upper part of the succession (L4 to L2). This species is
is known only by isolated vertebrae. It has been found in the Benguérir rather common in Level III (Upper Maastrichtian) of the Ouled Abdoun
Maastrichtian succession, except in L5 and “Sillon X”. It is more Basin (de Buffrenil et al., 2008; Houssaye et al., 2011).
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238
Fig. 9. Relative richness in number of collected teeth (total indicated at left) belonging to the largest predators at the various Maastrichtian phosphatic levels of Benguérir: Mosasauridae (left)
and Anacoracidae* (right). Abbreviations: M1: Globidens phosphaticus, M2: Prognathodon currii, M3: Prognathodon sp./Eremiasaurus heterodontus M4: Halisaurus arambourgi, M5: “Platecarpus”
ptychodon, M6: Mosasaurus beaugei, A1: Squalicorax pristodontus, A2: Squalicorax sp. 1, A3: Squalicorax sp. 2, A4: Squalicorax bassanii. *Squalicorax africanus is not considered here.
Mosasaurid squamates are represented by at least nine species and
are both abundant and diversified at all levels, especially in L3 and L2
(seven species).
Mosasaurus beaugei (Fig. 8H) is the rarest mosasaurid in the
Benguérir succession and is only represented in L2 by three in-
complete teeth. Teeth of this large mosasaurid (tooth crowns around
3 cm high) are more abundant in the upper Level III (upper
Maastrichtian) of the Ouled Abdoun Basin (Bardet et al., 2004). Its
occurrence both in L2 of Benguérir and in the upper Level III of the
Ouled Abdoun Basin suggests that its stratigraphical range is limited
to the Late Maastrichtian.
“Platecarpus” ptychodon (Fig. 8I) is a very characteristic small species
(tooth crowns around 1.5 cm high) whose teeth are very abundant, es-
pecially in L3. This species, attributed with doubt to the genus
Platecarpus by Arambourg (1952), is a plioplatecarpine russellosaurine
but belongs to a new genus.
Halisaurus arambourgi (Fig. 8J) is also a small species (tooth crowns
around 1 cm high) with very diagnostic teeth (Bardet et al., 2005a), well
represented in the succession except in “Sillon X”. It is particularly abun-
dant in L3 and L4, being by far the most abundant mosasaurid species in
L4 and the second in abundance in L3. Its abundance is roughly compa-
rable to that of “Platecarpus” ptychodon.
Globidens phosphaticus (Fig. 8M) is scarce (about 50 teeth) but is rep-
resented at all levels except L5 and “Sillon X”, with its highest abun-
dance in L4 and L3 (Bardet et al., 2005b).
Teeth of Carinodens belgicus (Fig. 8L) are very scarce elsewhere in the
phosphates of Morocco: up to now, only one has been found in L3 of
Benguérir, the other ones (including the tooth figured here) come
from Level III (Upper Maastrichtian) of the Ouled Abdoun Basin
(Bardet et al., 2008; Schulp et al., 2010).
The genus Prognathodon is the largest-sized mosasaurid of the as-
semblage (largest tooth crowns more than 5 cm high). It is the best rep-
resented taxon, being common in L6 and very abundant in L3 and L2. It is
the only marine reptile taxon identified in SX, where it remains however
very scarce. It is represented by three different species. P. giganteus
(Fig. 8O) is the commonest species from L6 and is absent in the upper
part of the succession. Its teeth are typically rounded in cross-section,
poorly posteriorly recurved and without pinched carinae. The tooth-
crowns of P. currii (Fig. 8N) are highly characteristic, being large high
and straight cones with a blunt apex (Bardet et al., 2005b). It is the
second rarest taxon after M. beaugei, only seven teeth having been
229
unearthed, but having been found however in all levels. Curiously,
the two largest teeth come from L5, the poorest level in vertebrate
remains of the Benguérir succession. A third species of Prognathodon,
here referred to as Prognathodon sp. (Fig. 8P), is the commonest one
in L3 and L2 but is absent in the lower part of the succession. The
teeth are similar to those of an undescribed new species from Level
III (Upper Maastrichtian) of the Ouled Abdoun Basin (N.B. unpub-
lished data).
Eremisasaurus heterodontus (Fig. 8K) is also a common species found
all through the Benguérir succession. It is mainly represented in L6 and
L4 to L2. This species has been described from Level III (Upper
Maastrichtian) of Grand Daoui area (Ouled Abdoun Basin) (LeBlanc
et al., 2012). Teeth here attributed to Prognathodon sp. and Eremiasaurus
heterodontus represent respectively the gracile, medium-sized teeth
and the robust, large ones described by Arambourg (1952) as
Mosasaurus cf. Leiodon anceps. Except for their different sizes, they are
difficult to differentiate, the probable reason why Arambourg referred
all of them to the same taxon.
To sum up, C. belgicus and M. beaugei appear to be restricted to the
upper part of the Maastrichtian of Benguérir. Considering their range
elsewhere, these species have probably a stratigraphical range restrict-
ed to the Late Maastrichtian and could thus be useful biostratigraphical
tools. On the other hand, the occurrence of P. giganteus (lower part of
the Benguérir section), versus Prognathodon sp. (upper part) reveals
that these two species could be also good biostratigraphical tools for dis-
crimination between the lower and the upper parts of the Benguérir
Maastrichtian succession.
Plesiosaurs are known by elasmosaurid isolated teeth (Fig. 8R)
and vertebrae. They are very common in L4 and L3 and common in
L6 and L2. Though these teeth and vertebrae are comparable to
both those of Plesiosaurus mauritanicus described by Arambourg
(1952), now considered a nomen dubium, and those of Zarafasaura
oceanis described by Vincent et al. (2011) from the Maastrichtian
of the Ouled Abdoun Basin, the new remains here collected exhibit
only general elasmosaurid characters and a more precise identifica-
tion is not currently possible.
Chelonians are represented throughout the succession by isolated
shell remains referable to both bothremydid pleurodirans and
chelonioid cryptodirans. They are more abundant in L6, L3 and L2.
Crocodyliforms are extremely scarce in the Benguérir succession.
Dyrosaurids have been recognised in L6, L4 and L2 on the basis of very
230 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238

few teeth and dermal scutes. Moreover, a single vertebra of an indeter- A 0 10 20 30 40 50

minate eusuchian is known from L6.
L2
7. Stratigraphical results 42.85

7.1. Stratigraphical level diversity and richness L3

6.25
Genera
7.1.1. General L4 excl. Genera
The effects on any environmental variations in foodweb complexity %
are usually highlighted by changes in top predator community. To as- L5
sess the quality of top predator community through the Maastrichthian,
we firstly calculated the relative richness in number of collected teeth
L6
belonging to eleven top predators throughout the different levels of 26.31
the Benguérir succession (Fig. 9). The top predators considered are the
mosasaurid squamates (seven taxa) and the anacoracid sharks (four
taxa, excluding S. africanus, which was a small-sized shark according B 0 10 20 30 40 50 60
to the size of its teeth). As far as the sharks are concerned, the domi-
nance of Squalicorax pristodontus compared to the other Squalicorax spe- L2
56.25
cies is noteworthy. Only in L6, where S. pristodontus is not so abundant
and represented by moderately large teeth, are the three other species L3
well represented. 10.52
Species
The same tendency is observed with the mosasaurids, with the
L4 excl. Species
dominance of Prognathodon sp./Eremiasaurus. The other species are 5.55
well represented only in L4 where Prognathodon sp./Eremiasaurus are %
less abundant. Prognathodon/Eremiasaurus represented the largest L5
mosasaurid predators, with probably a generalist diet compared to the
other species, which were generally smaller and had dental adaptations L6
indicating a more specialised diet (piscivorous for Halisaurus and 46.42

“P.” ptychodon, durophagous for Globidens). The scarcity of the large

generalist taxa P. currii and Mosasaurus is however notable and cannot C
be explained in the current state of the knowledge. 35
When mosasaurid and anacoracid occurrences are combined, a
30
general trend is apparent: anacoracids are dominant compared to
mosasaurids in the assemblages from L6 to L4. Their abundance is 25
rather comparable in L3. In L2, mosasaurids become much more
abundant compared to anacoracids. These richness fluctuations 20 Genera
could reveal a possible important trophic change difficult to explain Species
15
for the moment.
10
7.1.2. Selachians
Their remains are by far the most abundant vertebrate finds in all 5
levels of the Benguérir phosphatic succession. Table 1 gives the strati-
0
graphic range of the different taxa and the fluctuations of the assem- L6 L5 L4 L3 L2
blages are summarized in Fig. 10.
The number of genera and species varies strongly according to Fig. 10. Diversity of selachians genera (A) and species (B). The upper scale indicates the
the levels (Fig. 10C). The most diverse associations are observed in number of genera or species, and also the percentages of exclusive taxa. C: number of gen-
L6 and L2. The least diversified occurs in L5. Levels L4 and L3 show era and species by level.
an intermediate situation. In L4 and L3, the number of genera and
species increases, with an abrupt faunal diversification in L2.
Hexanchus and Pristiophorus, both occurring exclusively in L2, could assemblages than they really are. However, that has no conse-
be indicative of deeper waters. quence in terms of number of taxa at each level. All through the suc-
The stratigraphical range of genera and species shows important cession, Lamniformes and among these Squalicorax and Cretolamna,
variations throughout all the succession, particularly noteworthy in are particularly abundant.
L6 and L2 (Fig. 10A–C). In L6, more than 26% of the genera and From the base (L6) to the top (L2), one can observe important vari-
about 46% of the species are exclusive to this level. In L2, the corre- ations in the Lamniformes associations. The L6 and L2 levels show very
sponding values are about 43% for genera and more than 56% for spe- similar diversity of genera or species, but in the detail, the specific as-
cies. In L5, there are no exclusive genera or species. In L4, there are no semblages are very different between the two levels, with only three
exclusive genera and about 5% of exclusive species. Finally, in L3, species in common. The L6 unit contains the most diverse Lamniformes
about 6% of exclusive genera and about 10% of exclusive species assemblage, with 10 species belonging to four genera. The diversity de-
can be noted (Fig. 10). creases to six species and three genera in L5, as in L4 and L3, but in-
Only some Lamniformes, and particularly the genus Squalicorax, creases back up to five genera and 10 species in L2. In L5, the diversity
have been considered in terms of relative abundance through the drops down dramatically, possibly because of important changes in
succession. Most of the teeth of this order have been obtained by the environment. The Lamniformes are the dominant group and the mi-
surface collecting when the micro teeth have been obtained by croscopic teeth are scarce. The teeth of Squalicorax are much more nu-
sieving sediment. Therefore, there is an important bias in favour merous than are those of the other Lamniformes (Cretolamna and
of large teeth, which are much more abundant in the faunal Serratolamna), probably indicative of a more open marine environment
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238 231

(high sea level?) like what has been observed in the Maastrichtian of A relative abundance calculation for each mosasaurid species per
Angola (Antunes and Cappetta, 2002). stratigraphical level has also been performed (Fig. 12). Because the
teeth of Prognathodon nov. sp. and Eremiasaurus are sometimes difficult
to differentiate, they have been counted together. At most levels, the
7.1.3. Actinopterygians
most abundant species are always the large predators Prognathodon
All actinopterygians are present through the Maastrichtian Benguérir
and Eremiasaurus, as well as the ichthyophagous, medium- to small-
succession, at either the generic (Enchodus, Stratodus, Stephanodus) or
sized “Platecarpus” ptychodon and Halisaurus. It is interesting to note
familial (phyllodontids, pycnodontids) rank, except in “Sillon X”. These
that, when Prognathodon/Eremiasaurus are less abundant (L4 and L3),
assemblages appear thus very constant between the lower and the
the abundance of Halisaurus/“P.” ptychodon is higher, possibly indicative
upper parts of the Maastrichtian.
of a direct competitive exclusion or predation phenomenon among the
Among enchodontids however, interesting differential occurrences
mosasaurid assemblages through times. The cutting teeth of the large
have been observed between Enchodus elegans, that occurs from L6 to
Mosasaurus are very scarce, as are teeth of the highly specialised
L4 levels, and Enchodus libycus and Enchodus bursauxi, known from L4
durophagous Prognathodon currii (large predator), Globidens (medi-
to L2. These taxa appear thus to be useful biostratigraphical tools to dis-
um-size robust crushing form) and Carinodens (small and gracile
criminate between the lower and the upper part of the Maastrichtian
crushing form).
phosphatic succession of Benguérir.

7.1.4. Reptiles 7.2. Stratigraphic chart

After the selachians, their remains are the most abundant verte-
brates at all levels of the Benguérir phosphatic succession. Table 1
gives the stratigraphic range of the different taxa and the fluctuations
of the marine reptile and mosasaurid assemblages are respectively sum-
marized in Figs. 11 and 12.
The teeth and vertebrae of the most abundant and easy to identify
groups, the mosasaurid and pachyvaranid squamates as well as
the elasmosaurid plesiosaurs, have been counted all through the
Benguérir Maastrichtian levels. The total abundance of marine reptile
specimens (Fig. 11A) shows that the L3 unit is by far the richest level,
followed by L2, L4 and L6; L5 and “Sillon X” are the poorest. The number
of specimens has also been counted separately for each group and re-
veals that mosasaurid remains are by far the most abundant reptilian
find and constitute in each level of the Maastrichtian Benguérir succes-
sion always more than 2/3–3/4 of the data (Fig. 11B).
A show significant difference (Pdissimilarity b 10−2) due to the numerous
900
Number of specimens
Dissimilarity indices are useful tools for palaeoecologists. The Index
of Raup and Crick (Ir&c; Raup and Crick, 1979 as discussed in Harper,
1981) is a probabilistic index based on presence/absence data, which
is particularly powerful with palaeontological data (Travouillon et al.,
2007). Based on simulations (Legendre and Legendre, 1998), the non-
metric values of Ir&c are slightly below one when two communities
have no shared species and slightly above zero when two communities
are identical. By default, Ir&c were calculated between all levels, inde-
pendently from ages.
Such an analysis has been performed for all the selachian,
actinopterygian and marine reptile species occurring in the Benguérir
Maastrichtian succession (Fig. 13). For instance, species associations in
L6 and L5 seem undifferentiated (Pdissimilarity b 10−6) contrary to
the faunal associations recovered in L5 and L4 that could be considered
as easily separable (Pdissimilarity b 10−2). The faunal composition in L4
and L3 is statistically similar (Pdissimilarity b 10−11), showing very few
differences in species associations. On the contrary, levels L3 and L2
800 appearances of taxa, mainly selachians, in L2.

800 In order to relate the hierarchical degree of similarity between the

700 layers according to their faunal components, we performed an agglomer-
600 ative method (UPGMA) on the distance matrix Ir&c (Fig. 13). Without
461
500 423 temporal constraint and as expected from the stratigraphical distribution
400
261 of the species, two main groupings are highlighted, the lower one (L6–L5)
300
and the upper one (L4–L2) with Ir&c near 1 between L6 and L2, indicating
200
that the base and the top of the phosphatic succession contain two clearly
100 33
0 distinct species associations (with the exception of species known from
L6 L5 L4 L3 L2 base to top). The first association L6–L5 is considered as belonging to
Stratigraphical levels the Early Maastrichtian, with occurrences of Scapanorhynchus rapax,
Squalicorax africanus, Sq. nov. sp. 2, Cretolamna biauriculata, C. aff.
B maroccana, Rhombodus meridionalis, Duwibatis sp., all of them recovered
usually in well-dated Lower Maastrichtian beds elsewhere (for instance
Number of specimens in

100% Jordan, H.C. pers. obs.). The second association L4–L2 is ascribed to the
percentage of total

90%
80% Late Maastrichtian, with occurrences of Serratolamna serrata, S. africana,
70% Carcharias heathi, large teeth of Squalicorax pristodontus, Ixobatis
60%
50% mucronata and Rhombodus binkhorsti. Also of interest to note from these
40% analyses is that neither the actinopterygians nor the marine reptiles are
30%
20% informative as most taxa range throughout the Maastrichtian Benguérir
10% succession.
0%
L6 L5 L4 L3 L2 No geochronologic dating of the phosphatic series elsewhere in
Stratigraphical levels the world is currently available due to the special chemical conditions
of the phosphatic deposits. The completeness of the Maastrichtian in
Mosasauridae Pachyvaranidae Elasmosauridae
Benguérir remains unknown but we suspect two major gaps, consider-
ing the large faunal turnovers and richness variations shown by the
Fig. 11. Marine reptiles (squamates and plesiosaurs) abundance, as the number of teeth in
each phosphatic level (L6 to L2) of the Maastrichtian Benguérir series. A: Total specimen
palaeontological data. No environmental change affecting the marine
number. B: Specimen number for mosasaurid and pachyvaranid squamates as well as web was detected from predator associations, as suspected from the
elasmosaurid plesiosaurs expressed as a percentage of the total. lack of any evidence of sedimentological changes in the series.
232 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238

Mosasaurus beaugei
Globidens
1% "Platecarpus"
ptychodon
8. Palaeoecological results
phosphaticus 4%
0%
Prognathodon currii
Halisaurus arambourgi
8.1. Richness and faunal turnovers
0%
2%

Estimating of marine primary biomass (plankton, soft-bodied and

shelled invertebrates) is unavailable in the main phosphate ores due
Prognathodon sp. +
Eremiasaurus to the diagenetic dissolution of any carbonate shell material. Interpreta-
93%
tion of the palaeoecology in the Benguérir area is thus limited to the
predators possessing phosphatic hard tissues such as enamelled teeth.
A total of 82 species of marine top-predators have been recorded at
L2 Benguérir, with a maximum of 47 and 46 species recovered in L2 and
L6 respectively.
Globidens Temporal evolution of species richness for the various groups
phosphaticus
Prognathodon currii 5% Carinodens belgicus (batoid rays, sharks, actinopterygians and marine reptiles) is reported
0%
0% in Fig. 14 (dashed lines). It shows, from the base to the top of the succes-
"Platecarpus"
ptychodon sion, a rapid fall of diversity in L5 (mainly due to the scarcity of teeth in
25%
Prognathodon sp. + this level), followed by an increase until L2 for all groups except sela-
Eremiasaurus
39% chians, in which the diversity increase appears only in L3. The maximum
specific diversity for each group is in L6 and L2, where they are compa-
Halisaurus rable in absolute terms.
arambourgi
31% Appearance rate (appearances at the beginning of upper level/sum
of taxa in both levels) and extinction rate (disappearances at end of
the lower level/sum of taxa in both levels) were calculated for each suc-
L3 cessive pair of levels, by correcting the Lazarus effect (considering the
Lazarus taxa and thus, not the reappearances) or not. Results are also re-
Globidens ported in Fig. 14.
phosphaticus
Prognathodon currii
6% True extinction (no reappearance of Lazarus taxa) is highest for all
0%
"Platecarpus" groups between L6 and L5 (50% of extinction among batoids, about
ptychodon
24% 40% among sharks and 10% among marine reptiles) counterbalanced
Prognathodon sp. + by a similar renewal rate. In contrast, the L5/L4 boundary is marked
Eremiasaurus
26% by a slightly lower extinction rate (especially in sharks with disappear-
ance of 36% of species recovered in L5) counterbalanced by a higher re-
Halisaurus
newal rate (especially in batoids and actinopterygians). This fact
arambourgi
44%
explains why the faunal turnover is maximal between L5 and L4
(excepted for sharks where the diversity increase appears in L3 only)
and why the Early Maastrichtian (L6–L5) association is well differentiat-
L4 ed from those of the upper part of the Maastrichtian (L4–L2). Faunal
turnover is minimal between L4 and L3, with the lowest extinctions
and appearance rates. From L5–L4 and toward the top of the succession,
Prognathodon currii we observe a global increasing of renewal rate compared to extinction
7% "Platecarpus"
ptychodon rate (e.g. selachians).
22%
Use of uncorrected data (without correcting the Lazarus effect) can
change moderately the results (Fig. 14, horizontal lines), indicating
Prognathodon sp. + that some taxa successively disappear and reappear along the Benguérir
Eremiasaurus
45% succession, or that collecting effort is sometimes inconsistent. For ma-
Halisaurus
arambourgi rine reptiles, though the species richness remains in perfect agreement
26%
with that of the other vertebrate groups, differences are however nota-
ble for appearance and extinction rates, which are comparable all along
the succession, due to the fact that most species are present all along the
L5 Benguérir succession. Evolution of faunal turnover for actinopterygians
is currently unresolved, in part due to the inconsistence of the data
(dominated by supraspecific taxa).
Globidens

Prognathodon currii
phosphaticus Despite the apparent homogeneity of sedimentary deposits through
3%
0% the succession, the turnovers of predators are multiple. These may be
"Platecarpus"
ptychodon explained by a profound environmental change or major temporal
21%
gaps. No privileged assumption was retained between L6 and L5, as
Prognathodon sp. +
Eremiasaurus Halisaurus the material is too scarce in L5 (Fig. 14, dashed line). Extinction
61% arambourgi
15% and appearance rates are strongly well-balanced between L5 and
L4 with the highest appearance rate without evidence of sedimento-
logical change between these two marine deposits. Faunal turnover
is lowest at the L4/L3 boundary, suggesting little disturbance in fau-
nal composition between these two deposits. Faunal turnover is
L6
more evident between L3 and L2 and manifested by a high appear-
Fig. 12. Diversity and relative abundance of mosasaurid species through the Maastrichtian
ance rate (particularly in sharks), suggesting a change in food abun-
phosphatic levels (L6 to L2) of the Benguérir series. 0 per cent indicates those species very dance as testified by the rapid increase of marine predator diversity
poorly represented. (Fig. 14 — dashed line).
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238 233

Fig. 13. Left. Stratigraphic range of marine vertebrate taxa recovered in the Maastrichtian phosphatic series of Benguérir (sharks, reptiles, batoids and actinopterygians). Taxon names are
labelled and reported in Table 1. Right. Agglomerative analysis performed on dissimilarity index (Ir&c calculated on presence/absence of taxa between levels). Minimal values of Ir&c are
indicated on branch topology (*Lazarus taxa considered). See text for explanation.

6 8 10 12 14 16
6 8 10 12 14 16
L2
L2

BATOIDS
L3 L3 SHARKS

L4 L4

L5 L5

L6 L6

0 0,2 0,4 0,6 0,8 1 0 0,2 0,4 0,6 0,8 1

1 2 3 4 5 6 6 7 8 9 10 11
L2 L2

L3 L3

L4 L4

L5 BONY FISHES L5 REPTILES

L6 L6

0 0,2 0,4 0,6 0,8 1 0 0,2 0,4 0,6 0,8 1

Fig. 14. Evolution of species richness (number of species per level, upper scale respectively, dashed line) compared to the appearance (black box, lower scale) and extinction rates (white
box, lower scale) calculated between the successive Maastrichtian phosphatic levels of the Benguérir series (vertical scale), for batoids, sharks, actinopterygians and marine reptiles, re-
spectively. Horizontal lines indicate rate calculation without consideration of Lazarus taxa.
234 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238
8.2. Analysis of palaeoguilds
From an ecological point of view, a palaeoguild is defined as a group
of fossil species that probably used the same resource in a similar way,
space and time (DiMichele, 1994). Palaeoguilds are usually considered
in the fossil record, by regarding groups of animals sharing out a dental
similarity (Van Valkenburgh, 1995), time spans of 104 years or greater
being usual in palaeoecology. Palaeoguilds typically do not contain
several classes of vertebrates in order to avoid that the groups included
are not equally likely preserved (Lawton, 1984 in Van Valkenburgh,
1995). However, the phosphatic condition of fossilisation in Benguérir
seems similar for all the vertebrate groups considered (selachians,
actinopterygians and marine reptiles) only represented by isolated
teeth. To build our dental groups, we considered all these marine verte-
brates as marine top-predators. As it is true for recent species, the shape
and size of teeth belonging to fossil species may reflect their prey
preferences and thus reveal their palaeoecology. Massare (1987) al-
ready performed such analysis on Mesozoic marine reptiles (includ-
ing mosasaurids) and defined seven feeding preference guilds,
distributed between three main categories, including mollusc eaters
(“crush” guild), soft food eaters (“pierce” guild) and large-vertebrate
hunters (“cut” guild). Throughout the Jurassic, the array of guilds
present in the oceans worldwide remained stable although there
was a considerable taxonomic turnover within guilds. Unfortunately
Massare's guilds are inapplicable for selachian and actinopterygian
tooth morphology, leading us to propose a comparable but simpler
grouping as outlined in Fig. 15A. Three paleoguilds were detected
here from the tooth crown analysis: a first one characterized by a
grinding trend (“mollusc eaters”), a second one characterized by a
cutting trend (“large vertebrate hunters”) and a third one character-
ized by composite jaws.
The most similar is the tooth morphology (regarding their food con-
summation), the most close are the taxa in the dental/dietary
morphospace (Fig. 15A), independently of their phylogenetic affilia-
tions. Replaced in time (level by level, Fig. 15B), we can observe if
these dental/dietary morphospaces (and thus the trophic web) change
with time and, as a consequence, of the multiple faunal turnovers re-
ported before. As evidenced in Fig. 15B, only slight trophic web distur-
bances are observed through the levels, with only a short-term change
in L5. We can however notice that the animal group with grinding-
crushing teeth (“mollusc eaters”) tends to be less abundant and/or spe-
cialized in levels L5–L2 compared to L6. Similarly, animal group with
tearing–cutting teeth (“large vertebrate hunters”) seems to be less spe-
cialized in L4–L2 than in L6–L5, with a reinforcement to the top (L2) of
taxa sharing out teeth of tearing type. The animal group with composite
jaws seems more strongly affected by the short-term change in L5 but
their morphological diversity is recovered at the top of the succession
(L2).
The stability in food web is thus notable despite the large faunal
turnover rates between layers as noted before (Fig. 14). Besides a poten-
tial signal of an environmental change between L6 and L5 affecting the
predator community, few significant change in prey associations was
clearly detected.
It is noteworthy that the great increase of marine vertebrate
richness in L2 does not disturb the median dental/dietary
morphospace. This possibly indicates that there is no ecological di-
versification in L2 compared to L3 and older levels, and that only
the biomass increased in L2 and thus, the number of unspecialized
predators. This could explain why top predators such as large
mosasaurids tend to dominate large sharks progressively upsection
from the L4 unit, both in abundance and diversity aspects (Fig. 9 and
associated text). It is interesting to note that this tendency with in-
creasingly large apex predators toward the top of the Benguérir suc-
cession (i.e. the mosasaurid Prognathodon) is also seen in
chronomorphs of some of the dominant sharks (i.e. the lamniform
Squalicorax).
8.3. Palaeoecological and biostratigraphical comparisons with the Ouled
Abdoun Basin of Morocco
As far as marine reptiles are concerned and apart from some excep-
tions probably linked to their scarcity or restricted stratigraphical range,
most taxa are present in the Ganntour and the Ouled Abdoun phosphat-
ic basins of Morocco. This suggests that the ecological conditions in both
basins were probably comparable and not so different as previously
thought (Bardet, 2012). The fact that the Benguérir marine reptile taxa
occur throughout the Maastrichtian phosphatic succession also con-
firms that the comparable marine reptile assemblage of the Ouled
Abdoun Basin does not correspond to a mixture of faunas in relation
with a condensation of some levels of the phosphatic series but to a
real faunal association (Bardet, 2012).
The same can be concluded concerning the selachian assem-
blages. On the whole, the associations are very similar between the
Ganntour and Ouled Abdoun Basins. The main differences are
biostratigraphically related and are a consequence of the extreme
condensation of the series in the Daoui area, eastern part of the
Ouled Abdoun Basin. There, the equivalent of the L6 of Benguérir
could be observed only at the extreme base of the series, consisting
of less than 20 cm of a coarse phosphatic sediment rich in Roudereia
phosphatic molds, and broken and eroded teeth of Scapanorhynchus
rapax and Squalicorax nov. sp. 2. Currently, these levels, covered by
barren sediment during the mining operations, are no longer acces-
sible. For levels L4 to L2 of Benguérir, the faunal associations do not
show significant differences from those preserved in the Level III of
the Daoui area. In conclusion, we propose here biostratigraphical
correlations between the Benguérir Level 6 and the base of the
Daoui Level III, as well as between the Benguérir Levels 4 to 2 and
the Daoui Level III.
9. Palaeobiogeographical results
9.1. Selachians
Concerning selachians, several taxa seem to be restricted to the
southern Tethys margin. For instance, Squalicorax bassanii, S. africanus
and Squalicorax nov. sp. 2 are known in the Early Maastrichtian of
Egypt (Cappetta, 1991), Syria (Bardet et al., 2000) and Jordan
(Cappetta et al., 2000, and H.C. pers. obs.). Squalicorax nov. sp. 1 has
not been collected in Egypt nor in Jordan. Scapanorhynchus rapax is
also a typical Early Maastrichtian species, being abundant in the
Benguérir L6 unit, at the base of the Oued Erguita series (Souss Basin,
Morocco) and in Syria (Bardet et al., 2000). It is absent in the phosphatic
deposits of Jordan, but occurs here in older deposits, probably of Late
Campanian age (H.C., pers. obs.). Higher in the Maastrichtian succession,
this large predator disappears worldwide (H.C., pers. obs.). Serratolamna
khderii occurs in all Early Maastrichtian localities and is particularly
abundant in Jordan where it was first described (Zalmout and
Mustafa, 2001).
Among batoids, Hypsobatis weileri is very common in L6 whereas
Rhombodus meridionalis remains scarce, exactly the opposite to what
is observed at the base of the Maastrichtian succession of Oued Erguita.
The same observation can be made at the base of the Jordan and Syria
phosphatic succession, even if here these two species remain uncom-
mon. As all these previously mentioned localities can be considered as
equivalent in age, environmental differences can be invoked to explain
these faunal differences. It is important to note that all the characteristic
taxa of the Benguérir L6 unit are completely lacking in the northern Te-
thys margin.
9.2. Actinopterygians
As for selachians, several actinopterygian taxa of the Benguérir
Maastrichtian phosphatic succession seem to be restricted to the
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238 235

Fig. 15. Evolution of “dental/dietary morphospace” performed on dental similarity and size versus resource use, for marine vertebrate species recovered by level (as really observed) of the
Maastrichtian Benguérir phosphatic series. Tooth morphology categories are compiled for each species, sharing general size and gradual capacity for clutching–grinding or clutching–cut-
ting (see Table 1). More usefully illustrated in 2D using a Correspondence Analysis (A), each level was secondarily individualized (B) to reflect faunal association by time and sorted by
potential ability of predation (grouped here in clutching–grinding grade, clutching–cutting grade and composite jaw).

southern Tethys margin and could also be useful tools to define this
large palaeoprovince. This is the case with the three enchodontid spe-
cies described by Arambourg (1952), as well as with Stephanodus
libycus, which have been found in the Maastrichtian of Western and
North Africa, the Middle-East and South America (references in Bardet
et al., 2000), but remain unknown in coeval strata of the northern Te-
thys margin.
Other actinopterygian taxa of Benguérir have a widespread
palaeobiogeographical distribution, such as Stratodus apicalis, known
from the Campanian–Maastrichtian of North America, Western and
North Africa, and the Middle-East (references in Bardet et al., 2000).
These faunal differences have been interpreted as palaeoecological pref-
erences linked to palaeolatitudinal gradient differences (Bardet, 2012).
Endemic taxon is the mosasaurid Eremiasaurus heterodontus which
is known up to now only in the phosphates of Morocco (LeBlanc et al.,
2012). However, this species has been described very recently so that
a record bias cannot be excluded, new fieldworks could reveal the oc-
currence of this taxon elsewhere.
Cosmopolitan taxa include the mosasaurids Prognathodon giganteus,
known from the Campanian–Maastrichtian of Europe and Middle-East,
as well as Carinodens belgicus and Carinodens minalmamar found in the
Late Maastrichtian of the northern and southern Tethys margins (details
in Bardet, 2012).

9.3. Reptiles
More than half of the squamate association appears to be restricted
to the southern Tethys margin. Among them are the pachyvaranid
Pachyvaranus crassispondylus, found up to now only in the Maastrichtian
of Syria and Morocco (Houssaye et al., 2011), and the mosasaurids
Mosasaurus beaugei, “Platecarpus” ptychodon, Halisaurus arambourgi,
Globidens phosphaticus, Prognathodon currii and possibly Prognathodon
sp., all taxa found only in the Maastrichtian of the southern Tethys
Margin (Bardet, 2012). These squamates could thus be useful tools to
define this large palaeoprovince as they are quite different from those
of the northern Tethys and otherwise remain unknown elsewhere.
10. Conclusions
10.1. Palaeobiodiversity, turnovers and K/Pg crisis
The marine vertebrate palaeodiversity is high through all the
Maastrichtian Benguérir succession, with a decrease between L6 and
L5, mainly due to the scarcity of fossils at the latter. Moreover, none of
the groups seems to be declining. Even if the Danian levels have not
been considered in the present study, some remarks can be made
about the diversity trends of selachians, actinopterygians and marine
reptiles across the K/Pg boundary.
236 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238
Several of the selachian families occurring in the Maastrichtian do not
cross the K/Pg boundary (Cappetta, 1987). This is the case of the
Anacoracidae, Pseudocoracidae, Sclerorhynchidae and Rhombodontidae.
All are present and even diversified up to the top of Maastrichtian (L2).
For instance, 3 species of Squalicorax, 4 genera and species of
Sclerorhynchidae and abundant teeth of two species of Rhombodus
occur in L2. So, these groups were not declining just before they disap-
peared. The same can be noted for the genus Serratolamna, which is
abundant in L2.
While all actinopterygian families found in Benguérir have a large
stratigraphical range spanning the K/Pg boundary (and so being unin-
formative in this regard), this is not the case for the marine reptiles. In-
deed, pachyvaranid and mosasaurid squamates as well as elasmosaurid
plesiosaurs (other groups are currently poorly known) were wide-
spread and well diversified during the latest Cretaceous and became ex-
tinct during the K/Pg biological crisis. The Benguérir Maastrichtian
succession shows that, as in several other parts of the world, these ma-
rine reptiles were both abundant and diversified up to the top of the
stage. For example in L2, seven mosasaurid species (of the nine
recognised in Benguérir) are still present and abundant. As for sela-
chians, these groups were not on decline just before they disappeared.
10.2. Biostratigraphy
The great interest of phosphatic series is the great abundance and high
diversity of vertebrate remains they contain. Moreover, these remains –
most usually isolated teeth of selachians, actinopterygians, marine rep-
tiles, and more rarely mammals in Paleogene levels – are well preserved.
However, the reverse of the medal of this type of sediment is the scarcity
or even the complete lack of organisms with a calcareous skeleton like
mollusks (ammonites, bivalves, gastropods), foraminifera, or calcareous
nannofossils. Therefore, correlations with well calibrated vertebrate
faunas from series consisting in other types of sediments (chalk, lime-
stone) are quite difficult to establish.
Most of the well calibrated Maastrichtian marine vertebrate faunas
are known from northern Europe and North America. However, these
associations are rather different from those occurring in the south Te-
thyan realm making difficult their use for dating purposes. The most
complete Early and mid-Maastrichtian elasmobranch associations
from northern Europe have been published by Siverson (1993). Howev-
er, these associations from the Kristianstadt Basin (southern Sweden)
are very different from those observed in Moroccan phosphates and
do not allow useful comparison. For instance and concerning the sela-
chians, the Early Maastrichtian Swedish faunas contain diversified asso-
ciations of Orectolobiformes, Carcharhiniformes, Synechodontiformes
and Squaliformes, that is not the case in Morocco. The Lamniformes
are less diversified, with a complete lack of the genus Squalicorax. The
Swedish mid-Maastrichtian association is less diversified but shows
the same trends. Moreover, the Swedish localities have not yielded
Late Maastrichtian elasmobranch faunas.
The Upper Maastrichtian series of Maastricht contains elasmobranch
associations, which, unfortunately, have not been studied in detail, pre-
cluding a comparison with contemporaneous Moroccan faunas.
The Early Maastrichtian faunas from New Jersey (Cappetta and Case,
1975; Case and Cappetta, 2004) are particularly rich in Lamniformes,
with the genus Squalicorax including three species, S. pristodontus,
S. kaupi and S. sp. This last species is represented by a single tooth show-
ing a mesial cutting edge with a double serration like in the S. nov. sp. 2
from Benguérir, but with a tooth morphology close to that of S. kaupi.
Like in Moroco, a species of Scapanorhynchus, S. texanus, with teeth
smaller than those of S. rapax, is common (Cappetta and Case, 1975;
Case and Cappetta, 2004).
The Late Maastrichtian faunas of Texas (Case and Cappetta, 1997)
are rich and diversified, with 44 identified selachian species. However,
by their content, they clearly differ from the Moroccan faunas and a di-
rect comparison for dating purposes is hardly usable.
The age of the phosphates of Egypt (Duwi Formation) outcropping
along the Red Sea, between Quseir and Safaga, is dated as Early
Maastrichtian, mainly thanks to their dinokysts content (Schrank and
Perch-Nielsen, 1985; Cappetta, 1991). The base of the phosphate forma-
tion near Safaga, corresponding to the Early Maastrichtian, has yielded
teeth of Squalicorax africanus, a species which is also present at the
base of the phosphate series of Benguérir.
The phosphate deposits of Jordan occur from north to south of the
country, with the important ore of Eshidiya, in the southeast. The Late
Campanian or Early Maastrichtian age of these phosphates has been de-
bated (Abed and Amireh, 1999). They have been considered as Early
Maastrichtian on the basis of their foraminifera content (Hamam,
1977). As in the Lower Maastrichtian phosphate of the Red Sea, the elas-
mobranch association is characterized by the occurrence of Squalicorax
africanus.
In Syria, the phosphatic series is dated as Late Campanian/Early
Maastrichtian in age according to some authors (Atfeh, 1989; Bardet
et al., 2000). The base of the series in the Khneifiss deposits contains
an association with Squalicorax africanus, S. nov. sp. 2 (as Squalicorax
cf. S. yangaensis), Scapanorhynchus rapax, Rhombodus meridionalis,
Hypsobatis sp., indicative of an Early Maastrichtian age, compared to
the faunal association of Level 6 of Benguérir.
In Israel, phosphate deposits are well developed in the Oron syncline,
Negev desert. The elasmobranch fauna is rich and diversified from the
Upper Campanian to the Maastrichtian (Lewy and Cappetta, 1989).
The most important association comes from level 4a of Giva't Mador. Ac-
cording to the important change in the microfauna composition in this
level (Reiss, 1962) and coinciding with the first occurrence of
Aequipecten acuteplicatus Alth, 1850 of exclusive Maastrichtian age in
Europe (Dhondt, 1972), this level is considered as Early Maastrichtian
in age. However, the environmental conditions were very different
and the selachian fauna from Negev is difficult to compare directly
with those of Benguérir. The occurrence of deep water Squaliformes, of
numerous teeth of Hexanchiformes and of Pseudocorax are indicative of
bathyal conditions, contrary to neritic conditions prevailing in Benguérir.
A clear difference has been underscored between the lower part of
the Maastrichtian (L6–L5) with a selachian association very similar to
what can be observed in Syria and Jordan for instance (Scapanorhynchus
rapax, abundant Squalicorax bassanii, Serratolamna khderii, Hypsobatis
weileri, Parapalaeobates atlanticus) and its upper part (L4–L2) where
Cretolamna maroccana, Serratolamna serrata, large Rhombodus binkhorsti
are common. Almost all marine reptile species, mainly mosasaurids,
occur through the entire series, limiting their usefulness for Maastrichtian
biostratigraphy.
The distinct selachian associations in the lower and upper part of the
Maastrichtian are particularly important in terms of correlation and dat-
ing of the phosphatic succession in the southern Tethys realm. More-
over, the special morphology of the teeth of S. pristodontus occurring in
L2, compared to what is observed in lower levels, permits us to distin-
guish a lower Upper Maastrichtian and an upper Upper Maastrichtian
faunal assemblage.
From a biostratigraphical point of view, the Benguérir Maastrichtian
phosphatic succession is thus unique for both its completeness and high
marine vertebrate abundance and diversity, constituting a real “Rosetta
Stone”, with at least three different faunal assemblages allowing reliable
dating and correlations for most of the southern Tethys Maastrichtian
phosphatic levels.
10.3. Palaeoecology
The Moroccan phosphates have yielded rich and diversified marine
vertebrate faunas (Bardet et al., in press). These last decades, many
new genera and species have been described or revised, but despite
that, many new taxa remain to be described, mainly among selachians.
However, this situation is not an obstacle for using these taxa for
palaeoenvironmental or palaeoecological reconstructions. Indeed,
 H. Cappetta et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 409 (2014) 217–238
thanks to their dental types, defined independently of their system-
atic position (Cappetta, 1986c, 1987; Cappetta, 2012), the place of
selachians in the food chain and in the palaeoenvironment is easy
to deduct.
The succession of several phosphate levels through the Maastrichtian
Benguérir section is indicative of a relatively stable sedimentation envi-
ronment during a long period. Thanks to variations of the δ18O data
(Kocsis et al., 2014), changes of palaeotemperatures have been demon-
strated through the phosphate series. Nature of the sediment gives also
valuable information on the deposit conditions. For instance, the abun-
dance of coarse sand grains in Level 6 is indicative of a near shore
environment.
The environmental conditions, particularly depth, water temper-
ature, nature of the bottom substrate, etc., have a direct impact on
the vertebrate associations identified in the Moroccan phosphate de-
posits. In the Maastrichtian phosphate succession of Benguérir, geo-
chemical analyses on the apatite of selachian teeth have shown
variations from the base to the top of the series (Kocsis et al.,
2014). The negative shift of δ18O from L6 to L4 indicates an increase
of sea water temperature, and then a decrease from this level to the
top of the series. These variations are very similar to those observed
at Shatsky Rise, tropical Pacific (Jung et al., 2012). This warming ep-
isode has been noted in Poland and Ukraine on the basis of foraminif-
eral studies (Dubicka and Peryt, 2012) and seems to correspond with
an eustatic sea-level highstand.
The selachian diversity observed through the different levels of the
Benguérir succession shows the relative importance of the seven occur-
ring orders. Hexanchiformes and Pristiophoriformes are restricted to L2
with a species each representing 3% of the whole fauna.
In Lamniformes, the maximum rates are noted in L5 (50%), L6 (35%)
and L2 (30%). Their importance remains rather high in L4 and L3
(respectively 34% and 31%).
The Orectolobiformes show rather similar rates in L6 (10%), L2
(13%), L5 (6%), L4 (11%) and L3 (11%). The Carcharhiniformes are
represented in L6 and L2, with only a species in each level (3%). Re-
garding the batoids, Rajiformes and Myliobatiformes follow there-
abouts the same trends. The Rajiformes represents 35% of the L6
association, 25% of L5, 33% of L4, 32% of L3 and 29% of L2. As for
Myliobatiformes, 17% are present in L6, 19% in L5, 22% in L4, 26% in
L3 and 19% in L2.
The variations of the global diversity in the selachian associations
through the Benguérir phosphate succession can be correlated with
the environmental changes. For instance, the drop of Squalicorax
specific diversity between L5 and L4, with the extinction of
S. africanus and S. nov. sp. 2, could result from this warming episode
and/or from a sea-level highstand. It is also interesting to note that
the most diversified selachian associations occur in L6 and L2 corre-
sponding to cooler sea-water periods and sea-level lowstand. These
faunal trends are much less marked for marine reptiles and for
actinopterygians. However, it is important to keep in mind that
only large teeth corresponding to the largest actinopterygian genera
have been considered in this study.
No correlation has been observed between the increase of abun-
dance/diversity of predators and change in ecological niches. The abun-
dance of predators as well as a diversity increase not followed nor
correlated to an ecological niche increase has been evidenced through-
out the Benguérir Maastrichtian phosphatic succession. For instance, in
L2, the number of Squalicorax pristodontus teeth strongly decreases
while teeth of Cretolamna maroccana and Serratolamna serrata remain
abundant. This observation could be indicative of a prey composition
modification or of a competition with marine reptiles. Indeed, large
mosasaurids are here dominant, even if large sharks are still present,
which could explain the decline of anacoracids. Niche partitioning
among mosasaurid assemblages is also probably present as the smallest
species are more abundant in the levels where the largest are less and
vice versa.
237
10.4. Palaeobiogeography
Most taxa in the Benguérir Maastrichtian faunal associations are typ-
ical of the southern Tethys margin deposits (Egypt, Israel, Jordan, Syria).
Concerning selachians, Scapanorhynchus rapax, Cretolamna biauriculata,
Cretolamna maroccana, Cretolamna sp., Serratolamna kdherii, Squalicorax
bassanii, Squalicorax africanus, Squalicorax sp. 1, Squalicorax sp. 2,
Hypsobatis weileri, Rhombodus meridionalis, Coupatezia elevata, etc., can
be cited as the main examples. For actinopterygians, they are Enchodus
elegans, Enchodus bursauxi, Enchodus libycus and Stephanodus libycus. For
marine reptiles and more especially squamates, they are Pachyvaranus
crassispondylus, Mosasaurus beaugei, “Platecarpus” ptychodon, Halisaurus
arambourgi, Globidens phosphaticus, Prognathodon currii and possibly
Prognathodon sp.
It is noteworthy that selachian and mosasaurian associations in the
Benguérir Maastrichtian phosphatic succession are also found in other
countries of the southern Tethys margin, both in the Early Maastrichtian
(Middle East) and in the Late Maastrichtian, but remain absent in other
palaeoprovinces (northern Tethys margin for example). They could
thus be considered as characteristic of the whole Maastrichtian stage
of the southern Tethys margin. In the case of mosasaurids, these faunal
provinces have been linked to palaeoecological preferences related to
palaeolatitudinal gradients (Bardet, 2012).
Acknowledgements
The authors thank Dr. A. Schulp, an anonymous reviewer and the ed-
itor for their pertinent comments and suggestions. This work has
benefited from the scientific programme of collaboration between the
Office Chérifien des Phosphates (OCP, Casablanca, Morocco), the Ministère
de l'Energie, des Mines, de l'Eau et de l'Environnement (MEMEE, Rabat,
Morocco), the Muséum National d'Histoire Naturelle (MNHN, Paris,
France), and the Universities Cadi Ayyad (Marrakech, Morocco) and
Chouaîb Doukkali (El Jadida, Morocco).
The field work at Benguérir has always beneficiated from the sup-
port of the local OCP authorities. We thank Messrs Bazaoui and Miftah
for their assistance in the field. Financial support by the National Geo-
graphic (no. 6627-99), Action de Coopération CNRS/CNRST (no. 18567),
Programme International de Coopération Scientifique CNRS/CNRST (no.
4892), Programme ANR-PALASIAFRICA (ANR-08-JCJC-0017, ANR-ERC),
Ministerio de Economía y Competitividad of Spain (CGL2010-18851/
BTE) and Gobierno Vasco/Eusko Jaurlaritza (IT834-13), as well as by
funds from the Département Histoire de la Terre and UMR CNRS-
MNHN-UPMC 7207 (MNHN, Paris), and UMR CNRS 5554 (ISE-M, Mont-
pellier). ISE-M contribution no. 2014-061.
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