Geobios 37 (2004) 315–324

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Mosasaurus beaugei Arambourg, 1952 (Squamata, Mosasauridae)

from the Late Cretaceous phosphates of Morocco
Mosasaurus beaugei Arambourg, 1952 (Squamata, Mosasauridae)
du Crétacé supérieur des phosphates du Maroc
Nathalie Bardet a,*, Xabier Pereda Suberbiola a,b, Mohamed Iarochene c,
Fatima Bouyahyaoui c, Baadi Bouya d, Mbarek Amaghzaz d
a
Département histoire de la terre, UMR 5143 du CNRS, Muséum national d’histoire naturelle, 8, rue Buffon, 75005 Paris, France
b
Universidad del País Vasco/Euskal Herriko Unibertsitatea, Facultad de cienca y Tecnología, Departamento de Estratigrafía y Paleontología,
Apartado 644, 48080 Bilbao, Espagne
c
Ministère de l’énergie et des mines, direction de la géologie, BP 6208, Rabat, Maroc
d
Offıce chérifien des phosphates, centre minier de Khouribga, Khouribga, Maroc
Received 12 September 2002; accepted 18 February 2003
Available online 11 May 2004

Abstract
Mosasaurus beaugei Arambourg, 1952 was based on isolated teeth from the Maastrichtian phosphatic deposits of Morocco. The recent
discovery of new material, including skull and mandibular remains, improves our knowledge of this species. M. beaugei shares the following
synapomorphies with the genus Mosasaurus: large teeth bearing two prominent carinae and with asymmetrical labial and lingual surfaces, the
labial one being flattened and strongly facetted and the lingual one being convex; premaxillae with a small pointed rostrum and dentary without
rostrum; palatal elements closely united; coronoid with very large ventromedial process overlying the prearticular. M. beaugei is characterised
by the following autapomorphies: 12-13 maxillary teeth; marginal teeth bearing 3-5 prisms on the labial surface and 8-9 on the lingual one;
palatine with posterior border concave and perpendicular to the long axis of the skull; splenial visible laterally on half of the dentary ventral
surface; coronoid with anterior wing well developed and bearing two notches. M. beaugei is only known to date in the Maastrichtian
phosphates of Morocco.
© 2004 Elsevier SAS. All rights reserved.

Résumé
Mosasaurus beaugei Arambourg, 1952 est basée sur des dents isolées provenant des dépôts phosphatés du Maastrichtien du Maroc. La
découverte récente de nouveau matériel, y compris des restes crâniens et mandibulaires, permet d’améliorer notre connaissance de cette
espèce. M. beaugei partage les synapomorphies suivantes avec le genre Mosasaurus : grandes dents portant deux carènes proéminentes et des
surfaces labiale et linguale asymétriques, la surface labiale étant plate et fortement facettée, la surface linguale convexe ; prémaxillaire avec un
petit rostre pointu et dentaire sans rostre ; os du palais bien imbriqués entre eux ; coronoïde à grand processus ventromédial recouvrant le
préarticulaire. M. beaugei est caractérisée par les autapomorphies suivantes : 12-13 dents sur le maxillaire ; dents marginales portant 3-5
prismes sur la surface labiale et 8-9 sur la surface linguale ; palatin à bord postérieur concave et perpendiculaire à l’axe longitudinal du crâne ;
splénial visible latéralement sur la moitié de la surface ventrale du dentaire ; coronoïde avec une aile antérieure bien développée portant deux
entailles. M. beaugei est exclusivement connue dans les phosphates maastrichtiens du Maroc.
© 2004 Elsevier SAS. All rights reserved.

Keywords: Mosasaurus beaugei; Mosasauridae; Maastrichtian; Phosphates; Morocco

Mots clés : Mosasaurus beaugei ; Mosasauridae ; Maastrichtien ; Phosphates ; Maroc

* Corresponding author.
E-mail address: bardet@mnhn.fr (N. Bardet).

© 2004 Elsevier SAS. All rights reserved.

doi:10.1016/j.geobios.2003.02.006
316 N. Bardet et al. / Geobios 37 (2004) 315–324
1. Introduction
Due to an active collaboration between the Office Chéri-
fien des Phosphates (OCP), the Ministère de l’Énergie et des
Mines and the Centre National de la Recherche Scientifique
(CNRS), new palaeontological field work has been realised
in the Oulad Abdoun phosphatic Basin of Morocco over the
last few years. The results are the discovery of a great number
of marine vertebrate remains, especially mosasaurs, croco-
dilians and turtles. These new discoveries significantly im-
prove our knowledge of these marine reptiles, which were
previously known mainly by fragmentary or isolated remains
(Arambourg, 1952).
Mosasaurus beaugei was described by Arambourg (1952)
on the basis of few isolated teeth from the Maastrichtian
phosphatic deposits of Morocco. New material includes two
incomplete skulls with mandibles. These specimens are de-
scribed below along with their palaeobiogeographic implica-
tions.
Abbreviations: MNHN, Muséum National d’Histoire Na-
turelle, Paris, France; OCP-DEK/GE, Office Chérifien des
Phosphates-Khouribga, Service géologique, collection OCP,
Morocco.
2. Geographical and geological context
The phosphatic deposits of Morocco, known since 1908,
have been exploited as an economical resource since the
1920’s (Office Chérifien des Phosphates, 1989). They are
part of the Mediterranean Tethyan phosphogenic province,
which extends from North Africa to the Middle-East (Lucas
and Prévôt-Lucas, 1996). The phosphatic deposits of Mo-
rocco outcrop in four basins, which are from NE to SW:
Oulad Abdoun, Ganntour, Meskala, and Bu-Craa in the Sa-
hara (Fig. 1(A)). Stratigraphically, they extend from the Late
Cretaceous (Maastrichtian) to the middle Eocene (Lutetian),
spanning the largest interval of time of all Tethyan phos-
phates (Lucas and Prévôt-Lucas, 1996).
The phosphates of Morocco are very rich in marine verte-
brate remains, especially selachians, bony fishes and reptiles
(i.e. Arambourg, 1952; Noubhani and Cappetta, 1997; Cavin
et al., 2000). Recently, the Palaeogene and the Maastrichtian
deposits have yielded land mammals (see Gheerbrant et al.,
2001 and references), and pterosaur and dinosaur remains
(Pereda Suberbiola et al., 2003 and in press) respectively.
The Maastrichtian marine reptiles include plesiosaurs (elas-
mosaurids), squamates (mosasaurids and varanoids), turtles
(bothremydids and chelonioids) and very rare crocodilians.
Mosasaurids are the most abundant group. Based on Aram-
bourg (1952) and the new discoveries, the mosasaurid assem-
blage is composed of at least six species: Mosasaurus
beaugei, Platecarpus ? ptychodon, Prognathodon aurii,
Prognathodon sp.; Globidens sp. and a new species of Hali-
saurus (Bardet et al., in review).
Fig. 1. Map of Morocco showing the main phosphatic basins (A), location
map of Mosasaurus beaugei (Sidi Daoui zone) in the Oulad Abdoun Basin
(B), and stratigraphical log showing its occurrence into the phosphatic series
(C). Abbreviations: Li, limestones; Ma, marls; Ph, phosphates.
Fig. 1. Carte du Maroc montrant les principaux bassins à phosphate (A),
carte de localisation de Mosasaurus beaugei (zone de Sidi Daoui) dans le
Bassin des Oulad Abdoun (B), et colonne stratigraphique montrant sa
position dans la série phosphatée (C). Abréviations : Li, Calcaires ; Ma,
marnes ; Ph, Phosphates.
Mosasaurus beaugei is well represented by fossils in the
eastern part of the Oulad Abdoun phosphatic Basin, near the
city of Khouribga. Most of the remains come from the Sidi
Daoui zone of Grand Daoui, an actively quarried area for
phosphate (Fig. 1(B)). Sidi Daoui is by far the richest zone in
Maastrichtian marine vertebrates of the basin.
The Maastrichtian phosphatic series of the Oulad Abdoun
Basin (“Couche III” of the miners, or bed III) is very con-
densed, being only about 2-5 m thick. It is composed from
the bottom to the top of a basal grey limestone rich in fish
remains; yellow soft exploited phosphates (lower Couche
III); a thin yellow marly level; and grey with brown stripes
soft exploited phosphates (upper Couche III) (Fig. 1(C)).
Stratigraphically, the mosasaurid remains described here oc-
cur in the upper part of the Couche III, which is Late Maas-
trichtian in age on the basis of selachian teeth (Cappetta,
1987).
3. Systematic palaeontology
SQUAMATA Oppel, 1811
MOSASAURIDAE Gervais, 1853
MOSASAURINAE Gervais, 1853
Mosasaurus Conybeare, 1822
Type species: Mosasaurus hoffmanni Mantell, 1829
Diagnosis: see Lingham-Soliar (1995).
Mosasaurus beaugei Arambourg, 1952
Holotype: MNHN PMC 7, isolated tooth (Arambourg,
1952: pl. 39, Fig. 13).
Type locality and horizon: unknown locality, Oulad
Abdoun Basin, Morocco, Couche III; Late Cretaceous, Late
Maastrichtian.
 N. Bardet et al. / Geobios 37 (2004) 315–324
Referred specimens from the Oulad Abdoun Basin
(Couche III): MNHN PMC 8-9, two isolated teeth (Aram-
bourg, 1952: pl. 39, Figs. 14, 19), unknown locality; MNHN
PMC 10, isolated tooth (undescribed, Arambourg collec-
tion), André Delpit locality; MNHN PMC 11-13, three iso-
lated teeth (undescribed, Arambourg collection), Bou Jniba
locality; OCP-DEK/GE 83, incomplete skull and lower jaw,
Grand Daoui area, Sidi Daoui zone; OCP-DEK/GE 303,
incomplete skull and lower jaw, Grand Daoui area, Sidi
Daoui zone.
Diagnosis: large mosasaurid with 12-13 maxillary teeth;
marginal teeth bearing 3-5 prisms on the labial surface and
8-9 on the lingual surface; palatine with posterior border
concave and perpendicular to the long axis of the skull;
splenial visible laterally along half of the ventral dentary
length; coronoid with anterior wing well developed and bear-
ing two notches.
Comments on Arambourg collection: nine isolated
teeth from the Maastrichtian phosphates of both the Oulad
Abdoun and the Ganntour basins of Morocco were chosen as
the type series for Mosasaurus beaugei (Arambourg, 1952:
p. 282–284, pl. 39, Figs. 13–21). Among these, only three
teeth from the Oulad Abdoun Basin can be safely attributed
to this species: MNHN PMC 7 (holotype; see Arambourg,
1952: Fig. 13), PMC 8 (Fig. 14) and PMC 9 (Fig. 19). The
type specimen MNHN PMC 7 is an anterior tooth, bearing
3 labial prisms and indeterminate number of lingual ones.
MNHN PMC 8 is a tooth from the median part of the jaw and
bears 5 labial and 8 lingual prims. MNHN PMC 9 is a
posterior tooth, which bears 5 labial prisms and 8-9 lingual
ones.
Two teeth (Arambourg, 1952: Figs. 15, 21) come from the
Ganntour Basin. These teeth differ from M. beaugei in that
they are smaller and slender, more compressed laterally, with
a lenticular cross-section and bear 6 labial and lingual
prisms. Whether or not these differences are linked to sys-
tematics or ontogeny is currently unresolved. Consequently,
these teeth are currently not referred to M. beaugei. Other
four teeth described by Arambourg (1952: Figs. 16–18, 20)
do not belong to M. beaugei and can not be determined with
certainty.
Among the Arambourg collection, several unpublished
teeth from the Oulad Abdoun Basin can be referred to
M. beaugei (MNHN PMC 10-13). It should be noted that
M. beaugei is not abundant in the Oulad Abdoun Basin and
scarce in the Ganntour Basin, and only represented in this last
Basin in the top of the Maastrichtian series (“Couche 2”, Late
Maastrichtian).
4. Description
OCP-DEK/GE 303 (Fig. 2) is an incomplete skull with
mandible in left lateral view. As preserved, it is 103 cm long
and 50 cm high. The skull consists of the left maxilla, the
left jugal and fragments of the left temporal bar
317
(postorbitofrontal-squamosal), the suspensorium and, prob-
ably, the left quadrate. Fragments of the lateral ascending
branch of the suspensorium (prootic) contacting the su-
pratemporal are preserved ventral to the temporal bar. Just
ventral to the prootic and dorsal to the surangular are several
bone fragments that could belong to the left quadrate. Parts of
the pterygoid are preserved between the prootic and the
surangular, and just dorsal to the left splenial. The left man-
dibular ramus, visible in lateral view, is complete except for
the retroarticular process. Ventral to this ramus are the right
dentary, splenial and angular. Because of breakage the ante-
rior part of the right dentary is in medial view whereas the
posterior part, as well as the right splenial and angular, are in
lateral view. The right coronoid is fragmentary, only the
posterior part is preserved and displaced ventral to the right
splenial. In spite of crushing, the skull is articulated with the
mandible and the maxillary and dentary teeth remain in
occlusion, the jugal and the posterior process of the coronoid
lay close together, the lateral branch of the suspensorium
(prootic) is located ventral to the temporal bar, and the left
quadrate is in the glenoid cavity. The total skull length is
estimated to about 110 cm.
OCP-DEK/GE 83 (Figs. 3, 4) is a partial skull with man-
dible, preserved in two blocks. As preserved, the skull is
50 cm long from the tip of the snout to the posterior extremi-
ties of the maxillae and palatines (maximum width 27 cm).
The premaxilla and the crushed palatal bones lie in ventral
view, the left maxilla is preserved in ventrolateral view and
the right maxilla is in medial view. The two mandibular rami
are exposed in medial view (length about 74 cm), all bones
remaining in articulation. They are preserved from the tip of
the dentary to the middle part of the surangular (left ramus)
and to the glenoid cavity (right ramus). On the same block lay
the atlas-axis complex and the third cervical vertebra of a
plesiosaur. The estimated skull length of OCP-DEK/GE 83 is
about 80 cm.
For measurements see Table 1.
4.1. Skull
Premaxilla. It is V-shaped and bears two pairs of teeth.
There is a very small rostrum anterior to the teeth. The
internarial bar is not preserved.
Maxilla. It is very broad and deep, supporting 12-13 large
prismatic teeth. The terminal branches of the maxillary nerve
emerge from a row of large and nearby foramina located
dorsally to the lateral margin of the maxilla. On OCP-
DEK/GE 83, posteriorly the foramina are so close to each
other that they almost form a continuous groove. The tooth
row extends very far posteriorly, probably up to the middle
part of the horizontal ramus of the jugal. The medial and
lateral parapets are of equal high. The lateral suture with the
premaxilla is regularly oblique. The narial emargination is
not preserved.
Jugal. Both rami are slender and make a near right angle.
There is no evidence of a posteroventral process.
318 N. Bardet et al. / Geobios 37 (2004) 315–324

Fig. 2. Mosasaurus beaugei, OCP-DEK/GE 303, incomplete skull and lower jaw, Late Cretaceous (Maastrichtian), Oulad Abdoun Basin, Morocco. A,
photograph; B, interpretative drawing. Abbreviations: ar, articular; co, coronoid; lan, left angular; ld, left dentary; lj, left jugal; lsp, left splenial; mx, maxilla;
pofr, postorbitofrontal; pro; prootic; pt, pterygoid; Q, quadrate; ran, right angular; rd, right dentary; rj, right jugal; rsp, right splenial; san, surangular; sq,
squamosal V, vertebra.
Fig. 2. Mosasaurus beaugei, OCP-DEK/GE 303, crâne et mandibule incomplets, Crétacé supérieur (Maastrichtien), Bassin des Oulad Abdoun, Maroc. A,
photographie ; B, dessin interprétatif. Abréviations : ar, articulaire ; co, coronoïde ; lan, angulaire gauche ; ld, dentaire gauche ; lj, jugal gauche ; lsp, splénial
gauche ; mx, maxillaire ; pofr, postorbitofrontal ; pro, prootique ; pt, ptérygoïde ; Q, carré ; ran, angulaire droit ; rd, dentaire droit ; rj, jugal droit ; rsp, splénial
droit ; san, surangulaire ; sq, squamosal ; V, vertèbre.
 N. Bardet et al. / Geobios 37 (2004) 315–324 319

Fig. 3. Mosasaurus beaugei, OCP-DEK/GE 83, incomplete skull, Late Cretaceous (Maastrichtian), Oulad Abdoun Basin, Morocco. A, photograph; B,
interpretative drawing. Abbreviations: j, jugal; mx, maxilla; pa, palatine; pmx, premaxilla; vo, vomer.
Fig. 3. Mosasaurus beaugei, OCP-DEK/GE 83, crâne incomplet, Crétacé supérieur (Maastrichtien), Bassin des Oulad Abdoun, Maroc. A, photographie ; B,
dessin interprétatif. Abréviations : j, jugal ; mx, maxillaire ; pa, palatin ; pmx, prémaxillaire ; vo, vomer.

Postorbitofrontal-Squamosal. The temporal bar is thick
and robust. The oblique suture between the postorbitofrontal
and squamosal is at least 10 cm long.
Vomer-Palatine. As in Mosasaurus hoffmanni, the vomer
and palatine are tightly associated and the size of the open-
ings is reduced (Lingham-Soliar, 1995). The palatine firmly
contacts the maxilla laterally and the vomer medially. Its
posterior border is regularly concave and more perpendicular
to the longitudinal axis of the skull than in M. hoffmanni. The
vomer is long and slender, separated medially, and forms the
medial margin of the oval internal nares.
Pterygoid. At least six teeth or alveoli are preserved. The
teeth are small, posteriorly recurved, with a rounded cross-
section and finely ridged enamel.
320 N. Bardet et al. / Geobios 37 (2004) 315–324

Fig. 4. Mosasaurus beaugei, OCP-DEK/GE 83, subcomplete mandible, Late Cretaceous (Maastrichtian), Oulad Abdoun Basin, Morocco. A, photograph; B,
interpretative drawing. Abbreviations: an, angular; co, coronoid; d, dentary; prar, prearticular; san, surangular; sp, splenial.
Fig. 4. Mosasaurus beaugei, OCP-DEK/GE 83, mandibule subcomplète, Crétacé supérieur (Maastrichtien), Bassin des Oulad Abdoun, Maroc. A, photogra-
phie ; B, dessin interprétatif. Abréviations : an, angulaire ; co, coronoïde ; d, dentaire ; prar, préarticulaire ; san, surangulaire ; sp, splénial.
 N. Bardet et al. / Geobios 37 (2004) 315–324 321

Table 1 with a median groove. The medial ascending wing is very

Measurements (in cm) of the specimens OCP-DEK/GE 83 and 303 from the high and straight, almost reaching the dorsal margin of the
Late Maastrichtian of the Oulad Abdoun Basin of Morocco.
Mesures en cm des spécimens OCP-DEK/GE 83 et 303 du Maastrichtien
dentary and the anterior margin of the coronoid.
supérieur du Bassin des Oulad Abdoun du Maroc Angular. The articulation for the splenial is oval, convex,
and bears a median vertical waved crest. Both the lateral and
OCP-DEK/GE OCP-DEK/GE
83 303
medial wings appear to be low.
Maxilla Length 45 cm 61 cm
Surangular. This is a large rectangular bone, as usual in
Height 9cm (9th tooth) 10 cm (6th tooth) mosasaurids. The anterior margin is obliquely straight. The
Jugal Length (horiz. - 20 cm lateral exposure of the coronoid surface is large and strongly
ramus) ridged. The surangular is well exposed between the coronoid
Height (horiz. - 2 cm and the angular on the lateral side of the mandible. Posterior
ramus) to the coronoid, the surangular descends strongly to the
Length (vert. ramus) - 6.5 cm glenoid fossa. Its dorsal margin is slightly curved, convex
Height (vert. ramus) - 3 cm anteriorly and concave posteriorly. The surangular takes part
Temporal bar Length - more than 12 cm
in the anterior and lateral borders of the glenoid fossa, the rest
Width - 3 cm
of the fossa being formed by the articular. The ventral suture
Prootic Length - 7 cm
Height - 3.5-4 cm
with the angular and articular is straight and almost horizon-
Dentary Length 49 cm 65 cm tal.
Anterior Height 3 cm 4.5 cm Coronoid. This is a very large bone with a general form
Posterior Height 9-9.5 cm 10 cm very similar to that of Prognathodon overtoni and to other
Splenial Length 45 cm more than 40 cm species of Mosasaurus (see Russell, 1967). In lateral view,
Medial wing Height 9 cm - the anterior process is a large oblique wing bearing two
Angular Length - 15 cm (left) notches. The lateral process is regularly convex. The medial
Medial wing Height 6 cm - process is very well developed and extends almost to the
Articulation Height 3 cm 4 cm angular. The posterior dorsal process is large and bears ridges
Surangular Length more than more than 35 cm on its medial surface.
23 cm
Articular-Prearticular. The prearticular is projected an-
Post. Coronoid Hei- 7 cm or more 13.5 cm
ght teriorly through the intramandibular join between the dentary
Ante. Glenoid Hei- - 9 cm and splenial. The articular occupies most of the glenoid
ght fossa, which is roughly oval and slightly concave. The ret-
Coronoid Length 13.5-14 cm 20 cm roarticular process is broken.
Medial Height 7 cm (medial) 7 cm (lateral)
Total Medial Height 11 cm (medial) 12.6 cm (lateral) 4.3. Marginal dentition
Teeth Crown Height 1.8-3.5 cm 5 cm
The teeth of the two specimens are similar to MNHN
Basal crown Length 1.5-2 cm 2.5 cm
PMC 7-9 described by Arambourg (1952) for Mosasaurus
beaugei. The dentition is homogenous along the jaws al-
The suspensorium (prootic) and quadrate are too fragmen- though the posterior teeth are slightly lower and wider than
tary for an accurate description. the others and the maxillary teeth are slightly larger than the
dentary ones. The root of the teeth is very large and barrel-
4.2. Lower jaw shaped. The robust crowns are high and relatively slender
(height about twice the basal width), slightly posteriorly
Dentary. The dentary terminates abruptly in front of the recurved. They bear two carinae with minute crenulations.
first tooth. It bears 14 or 15 teeth on OCP-DEK/GE 83 and The labial surface is much smaller than the lingual one, with
16 on OCP-DEK/GE 303. The depth of the dentary increases a U-shaped cross-section as typical in the genus Mosasaurus
regularly from the anterior tip to the posterior margin. Two (Russell, 1967). This cross-section is much marked on the
rows of foramina are present on the anterior third of the anterior teeth than in the posterior ones, which exhibit a more
lateral surface. The dorsal and ventral margins are straight. lenticular compressed form. The labial surface bears between
The medial and lateral parapets are of equal height. The 3 (anteriormost teeth) and 5 prisms, being 4-5 the most
retrodental process is very short. The Meckelian canal ex- common. The medial surface bears about 8-9 prisms, which
tends almost to the anterior end. On OCP-DEK/GE 303, the generally do not reach the apex. The prisms are less discern-
posterior border of the dentary bears a dorsal horizontal able on the posteriormost teeth of the jaws and on largest
excavation like in Prognathodon overtoni (Russell, 1967). specimens.
Splenial. It extends medially over two thirds of the den-
tary length and on the lateral surface it extends along half the
5. Discussion
ventral edge of the dentary. The posterior portion is robust,
expanding posteriorly to the dentary and bears ventrally a The specimens OCP-DEK/GE 83 and 303 show two syna-
large oval foramen. The articulation for the angular is oval pomorphies of the Mosasaurinae: a short conical premaxil-
322 N. Bardet et al. / Geobios 37 (2004) 315–324
Table 2
Comparison of maxillary and dentary tooth number and ornamentation in Mosasaurus species.
Comparaison du nombre de dents et ornementation chez les espèces de Mosasaurus
beaugei hoffmanni
Maxilla 12–13 14
Dentary 14-16 14
Tooth size large large
Labial prisms 3–5 2–3
Lingual prisms 8–9 no
lary rostrum (DeBraga and Carroll, 1993: character 4a; Bell,
1997: character 2-1), and a surangular with a large ascending
coronoid buttress (DeBraga and Carroll, 1993: 57a; Bell,
1997: 78-1). The taxonomic composition of this clade varies
depending on the authors: for DeBraga and Carroll (1993) it
consists of (Clidastes + (Globidens + (Mosasaurus + Ploto-
saurus))) whereas for Bell (1997) it includes (Clidastes +
(Globidens + (Prognathodon + Plesiotylosaurus)) + (Mosa-
saurus + Plotosaurus))). In the following discussion, the
specimens from Morocco will be compared with mosasau-
rine taxa in its largest sence (sensu Bell, 1997).
The Morocco specimens share with all mosasaurines but
Clidastes (Bell, 1997: node I, Globidensini + Plotosaurini) a
coronoid with a very developed medial wing that nearly
contacts or contacts the angular (DeBraga and Carroll, 1993:
60b-c; Bell, 1997: 76-1). Moreover, OCP-DEK/GE 83 and
303 greatly differ from Clidastes (type species C. propython)
in having no dentary rostrum and in the shape and number of
both maxillary and dentary teeth.
The Morocco specimens differ from all the Globidensini
of Bell (1997) in the presence of high teeth with asymmetri-
cal labial and lingual surfaces. Moreover, they differ from
Globidens in having pterygoid teeth and a coronoid with a
posterior process poorly ossified to the surangular; and from
Prognathodon and Plesiotylosaurus in having pterygoid
teeth smaller than the marginal ones and a straight dentary
ventral border (DeBraga and Carroll, 1993; Bell, 1997). They
differs from Plesiotylosaurus in having no dentary rostrum
and a splenial that does not reach the symphysis; and from
Prognathodon in having a large coronoid medial process that
does not contact the angular.
With regard to the Plotosaurini clade, only a synapomor-
phy can be checked on the Morocco specimens: a straight
suture between the surangular and angular (Bell, 1997). Plo-
tosaurus differs from OCP-DEK/GE 83 and 303 in having a
larger number of both maxillary and dentary teeth, a long
dentary rostrum, and an angular covering most of the medial
surface of the articular (Russell, 1967; DeBraga and Carroll,
1993; Bell, 1997).
OCP-DEK/GE 83 and 303 can be safely assigned to the
genus Mosasaurus on the basis of the following synapomor-
phies: teeth greatly enlarged bearing two prominent carinae
and with asymmetrical buccal and lingual surfaces, the buc-
cal one being flattened and strongly facetted and the lingual
one being convex (DeBraga and Carroll, 1993: 52b; Russell,
1967; Lingham-Soliar, 1995); 13-15 maxillary and 14-15
dentary teeth (Russell, 1967); premaxilla with a small
maximus conodon lemonnieri missouriensis
14 ? 15 14
14 17 17 14-15
large ? slender large
2–3 ? 8–10 4–6
no ? ? 8
pointed rostrum and dentary without any rostrum (Russell,
1967); palatal elements closely united (Lingham-Soliar,
1995); coronoid with very large ventromedially process
overlying prearticular (DeBraga and Carroll, 1993: 60b,
Lingham-Soliar, 1995). Lingham-Soliar (1995) has consid-
ered the posterior extension of maxillary teeth below the
orbit as apomorphic for Mosasaurus but this character is
regarded as plesiomorphic by DeBraga and Carroll (1993:
character 11).
Currently, no revision of Mosasaurus has been attempted,
the only available descriptions are those of the North Ameri-
can species by Russell (1967) and of M. hoffmanni by
Lingham-Soliar (1995) and Mulder (1999) and of M. lemon-
nieri by Lingham-Sdiar (2000). According to these authors,
Mosasaurus includes several species, most of them from
Europe and North America. In Europe, M. hoffmanni Man-
tell, 1829 (the first mosasaurid that was found and named)
and M. lemonnieri Dollo, 1889, both from the Maastrichtian
of The Netherlands and Belgium are known. In North
America, there are the following species (Russell, 1967):
M. missouriensis (Harlan, 1834) from the Campanian of
Missouri and South Dakota; M. dekayi Bronn, 1838 from the
Maastrichtian of New-Jersey; M. maximus Cope, 1869 from
the Maastrichtian of New-Jersey and Texas; and M. conodon
(Cope, 1881) from the Campanian of Arkansas and South
Dakota and the Maastrichtian of New Jersey. Recently,
M. ivoensis Persson, 1963, from the Campanian of Sweden
and Kansas, has been reassigned to Tylosaurus (Lindgren
and Siverson, 2002). Baird and Case (1966) and Russell
(1967) regarded provisionally M. lemonnieri as a junior
synonym of M. conodon. According to Mulder (1999),
M. dekayi and M. maximus are junior synonyms of M. hoff-
manni. However, Lingham-Soliar (1995) considered both
M. lemonnieri and M. maximus as valid species. In his
phylogenetical analysis of North America mosasaurids, Bell
(1997) includes only M. conodon, M. maximus and M. mis-
souriensis as valid species of Mosasaurus. Pending a revi-
sion of this genus, the Morocco specimens have been com-
pared to M. hoffmanni-maximus, M. conodon-lemonnieri
and M. missouriensis. Comparisons with M. dekayi, based
on an isolated tooth (Russell, 1967), have not been consid-
ered.
Mosasaurus beaugei differs mainly from other Mosasau-
rus species in the number of maxillary teeth and the number
of prisms on the enamel crown (see Table 2). Indeed,
M. beaugei exhibits a lower number of maxillary teeth (only
12-13 pairs) than any other Mosasaurus species (all of them
 N. Bardet et al. / Geobios 37 (2004) 315–324
having 14-15). With regard to the crown ornamentation,
M. beaugei shows 3-5 labial prisms, which differs clearly
from M. hoffmanni-maximus (2-3 prims), and M. conodon-
lemonnieri (8-10). M. missouriensis has a similar number of
labial (4-6) and lingual (8) prisms but differs from
M. beaugei in having more teeth on the maxilla. The number
of dentary teeth, which varies between 14 and 16 pairs in
M. beaugei, ranges in the interval of Mosasaurus. M.
beaugei is also characterised by the following features: pa-
latine perpendicular to the long axis of the skull (oblique in
M. hoffmanni-maximus); splenial visible laterally on half of
the ventral dentary surface (only the posterior third in M.
hoffmanni-maximus); and coronoid with well developed an-
terior wing bearing two notches (less developed and bearing
a single notch in M. hoffmanni-maximus; less developed in
M. lemonnieri and M. missouriensis).
6. Palaeobiogeographical implications
Mosasaurus beaugei is only known to date from the Late
Maastrichtian phosphates of Morocco. Bardet and Pereda
Suberbiola (1998) have suggested that differences in the
mosasaurid assemblages from the Maastrichtian of the north-
ern and southern margins of the Mediterranean Tethys could
be linked to paleolatitudinal gradients, as already suggested
by Nicholls and Russell (1990) for the Late Cretaceous
marine vertebrate assemblages from the Western Interior
Seaway of North America. Considering the above-mentioned
synonymies, two species of Mosasaurus are known in the
Late Maastrichtian of Europe and North America, i.e. M.
hoffmanni-maximus and M. conodon-lemonnieri (Russell,
1967; Mulder, 1999). M. hoffmanni has also been described
in the Late Maastrichtian of Turkey (Bardet and TunoWlu,
2002). This suggests that the distribution of these species of
Mosasaurus is restricted to a paleolatitudinal belt around 30°
N. In the southern margin of the Mediterranean Tethys
(northern Africa), only M. beaugei is represented in the Late
Maastrichtian. The conspicuous absence of this species in the
phosphates of Egypt and Middle-East could be related to the
older age of these deposits, which are probably Early Maas-
trichtian (see Bardet et al., 2000 for more details). If correct,
this palaeobiogeographical hypothesis should be confirmed
by the discovery of M. beaugei remains in Late Maastrich-
tian levels located at around 20° N.
Acknowledgements
This work has benefited from the help and collaboration
(“Tripartite Convention”) of the Direction de la Géologie
from the Ministère de l’Énergie et des Mines (Rabat) and of
the Office Chérifien des Phosphates (Casablanca) from the
Kingdom of Morocco. The senior authors especially thank
MM M. Hamdi, M. Zeghnoun, and all the staff of the OCP
mining center for their active support during their stay in
323
Khouribga as well as MMs M. Sadiqui, L. Tabit and N.
Aquesbi from the Ministère de l’Énergie et des Mines (Ra-
bat) for providing administrative facilities and permits. Pho-
tographs are by MM D. Serrette and P. Loubry (CNRS,
MNHN, Paris). Infography of the figures is by H. Lavina
(CNRS, MNHN, Paris). This research work was supported
by funds from the CNRS and the National Geographic Soci-
ety (Grant #6627-99). The authors warmly thank the two
referees E.L. Nicholls and E.W.A. Mulder for their helpful
and constructive comments.
References
Arambourg, C., 1952. Les vertébrés fossiles des gisements de phosphates
(Maroc – Algérie – Tunisie). Notes et Mémoires du Service géologique
du Maroc 92, 1–372.
Baird, D., Case, G.R., 1966. Rare marine reptiles from the Cretaceous of
New Jersey. Journal of Paleontology 40, 1211–1215.
Bardet, N., Cappetta, H., Pereda Suberbiola, X., Mouty, M., Al Maleh, A.K.,
Ahmad, A.M., Khrata, O., Gannoum, N., 2000. The marine vertebrate
faunas from the Late Cretaceous phosphates of Syria. Geological Maga-
zine 137, 269–290.
Bardet, N., Pereda Suberbiola, X., 1998. Distribution patterns of Late Cre-
taceous marine reptile faunas from the Tethyan margins: palaeobiogeo-
graphical implications. In: Gayet, M., Otero, O. (Eds.), Palaeodiversifi-
cations land and sea compared, Lyon.
Bardet, N., TunoWlu, C., 2002. The first mosasaur (Squamata) from the Late
Cretaceous of Turkey. Journal of Vertebrate Paleontology 22, 712–715.
Bell, G.B., 1997. A phylogenetical revision of North American and Adriatic
Mosasauroidea. In: Callaway, J.M., Nicholls, E.L. (Eds.), Ancient
Marine Reptiles. Academic Press, San Diego, pp. 293–332.
Bronn, H.G., 1838. Lethaea geognostica. Stuttgart 2, 545–1346.
Cappetta, H., 1987. Mesozoic and Cenozoic Elasmobranchii, Chondrich-
thyes II. In: Schultze, H.-P. (Ed.), Handbook of Paleoichthyology.
Gustav Fischer Verlag, Stuttgart and New York, vol. 3B, 193 p.
Cavin, L., Bardet, N., Cappetta, H., Gheerbrant, E., Iarochène, S.M.,
Sudre, J., 2000. A new Palaeocene albulid (Teleostei: Elopomorpha)
from the Ouled Abdoun phosphatic basin, Morocco. Geological Maga-
zine 137, 583–591.
Conybeare, W.D., 1822. In: Parkinson, J. (Ed.), Outlines in Oryctology: an
introduction to the study of fossil organic remains 1st ed. London.
Cope, H.D., 1869. On the reptilian orders Pythonomorpha and Streptosau-
ria. Boston Society of Natural History Proceedings 12, 250–266.
Cope, H.D., 1881. A new Clidastes from New Jersey. American Naturalist
15, 587–588.
DeBraga, M., Carroll, R.L., 1993. The origin of mosasaurs as a model of
macroevolutionary patterns and processes. In: Hecht, M.K., et al. (Eds.),
Evolutionary Biology, 27. Plenum Press, New York, pp. 245–304.
Dollo, L., 1889. Première note sur les mosasauriens de Mesvin. Société
Belge de Géologie et de Minéralogie 3, 271–304.
Gervais, P., 1853. Observations relatives aux reptiles fossiles de France.
Comptes Rendus de l’Académie des Sciences de Paris 36, 374–377 et
470–474.
Gheerbrant, E., Sudre, J., Iarochène, M., Moumni, A., 2001. First ascer-
tained African “condylarth” mammals (primitive ungulates: cf. Bulbulo-
dentata and cf. Phenacodonta) from the earliest Ypresian of the Ouled
Abdoun Basin, Morocco. Journal of Vertebrate Paleontology 21, 107–
118.
Harlan, R., 1834. On some new species of fossil saurians found in America.
British Association for the Advancement of Sciences 1833, 440.
Lindgren, J., Siverson, M., 2002. Tylosaurus ivoensis: a giant mosasaur from
the early Campanian of Sweden. Transactions of the Royal Society of
Edinburgh, Earth Sciences 93, 73–93.
324 N. Bardet et al. / Geobios 37 (2004) 315–324
Lingham-Soliar, T., 1995. Anatomy and functional morphology of the larg-
est marine reptile known, Mosasaurus hoffmanni (Mosasauridae,
Reptilia) from the Upper Cretaceous, Upper Maastrichtian of The Neth-
erlands. Philosophical Transactions of the Royal Society of London
B347, 155–180.
Lucas, J., Prévôt-Lucas, L., 1996. Tethyan phosphates and bioproductites.
In: Nairn, A.E.M., et al. (Eds.), The Ocean Basins and Margins. Plenum
Press, New-York, pp. 367–391.
Mantell, G.A., 1829. A tabular arrangement of the organic remains of the
county of Sussex. Geological Society of London Transactions 3, 201–216.
Mulder, E.W.A., 1999. Transatlantic latest Cretaceous mosasaurs (Reptilia,
Lacertilia) from the Maastrichtian type area and New Jersey. Geologie en
Mijnbouw 78, 281–300.
Nicholls, E.L., Russell, A.P., 1990. Paleobiogeography of the Cretaceous
Western Interior Seaway of North America: the vertebrate evidence.
Palaeogeography, Palaeoclimatology, Palaeoecology 79, 149–169.
Noubhani, A., Cappetta, H., 1997. Les Orectolobiformes, Carcharhiniformes
et Myliobatiformes (Elasmobranchii, Neoselachii) des bassins à phos-
phate du Maroc (Maastrichtien-Lutétien basal). Systématique, biostratig-
raphie, évolution et dynamique des faunes. PalaeoIchthyologica 8, 1–327.
Office Chérifien des Phosphates, 1989. The phosphates basins of Morocco.
In: Notholt, A.J.G., Sheldon, R.P., Davidson, D.F. (Eds.), Phosphates
deposits of the world, volume 2 – Phosphate rock resources. Cambridge
University Press, Cambridge, pp. 301–311.
Oppel, M., 1811. Die Ordnung + Familien und Gattungen der Reptilien als
Prodrom einer Naturgeschichte derselben. Münich.
Pereda Suberbiola, X., Bardet, N., Jouve, S., Iarochène, M., Bouya, B.,
Amaghzaz, M., in press. A new azhdarchid pterosaur from the Late
Cretaceous phosphates of Morocco. Journal of the Geological Society of
London, Special Publication, 217: 79–90.
Persson, P.O., 1963. Studies on Mesozoic marine reptile faunas with particu-
lar regard to the Plesiosauria. Inst. Min. Paleont Quaternary Geology of
the University of Lund Publication 118, 1–15.
Russell, D.A., 1967. Systematics and morphology of American mosasaurs.
Bulletin of the Peabody Museum of Natural History 23, 1–240.
Williston, S.W., 1897. Range and distribution of the mosasaurs. Kansas
University Quarterly 6, 177–189.