Next Article in Journal
Minor Title Change: Fossils Becomes Fossil Studies
Previous Article in Journal
The First Dinosaur from the Kingdom of Cambodia: A Sauropod Fibula from the Lower Cretaceous of Koh Kong Province, South-Western Cambodia
 
 
Search for Articles:
Title / Keyword
Author / Affiliation / Email
Journal
Article Type
 
 
Advanced Search
 
Section
Special Issue
Volume
Issue
Number
Page
 
Journals
Fossil Studies
Volume 1
Issue 1
10.3390/fossils1010007
fossstud-logo
Submit to this Journal Review for this Journal Propose a Special Issue
► Article Menu

Article Menu

share Share announcement Help format_quote Cite question_answer Discuss in SciProfiles
first_page
settings
Order Article Reprints
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Background:
Article

A Prognathodontin Mosasaur from the Maastrichtian of the Dakhla Oasis, Western Desert, Egypt

by
Gebely A. Abu El-Kheir
1,
Ahmed A. Shaker
2,
Hallie P. Street
3,
Nicholas R. Longrich
4,*,
Amin Strougo
2,
Anhar Asan
2 and
Mohamed AbdelGawad
5
1
New Valley Vertebrate Palaeontology Centre, New Valley University, Kharga City 172201, Egypt
2
Department of Geology, Faculty of Science, Ain Shams University, Cairo 11566, Egypt
3
Department of Biological Sciences, MacEwan University, Edmonton, AB T5J 4S2, Canada
4
Department of Biology and Biochemistry, University of Bath, Bath BA2 7AY, UK
5
Department of Geology, Faculty of Science, Cairo University, Giza 12613, Egypt
*
Author to whom correspondence should be addressed.
Foss. Stud. 2023, 1(1), 60-75; https://doi.org/10.3390/fossils1010007
Submission received: 28 July 2023 / Revised: 29 September 2023 / Accepted: 6 October 2023 / Published: 9 November 2023
Browse Figures
Versions Notes

Abstract

:
Mosasaurs were diverse in the Upper Cretaceous in Africa, but relatively little is known about the mosasaur fauna of Egypt. Here, associated teeth and postcranial skeletal elements are reported for a mosasaur from the Maastrichtian Dakhla Shale of the Dakhla Oasis. The specimen includes tooth crowns, cervical, dorsal, and caudal vertebrae, and ribs. Teeth and bones exhibit features allowing referral to Prognathodontini. The teeth are relatively straight and blunt, suggesting affinities with Prognathodon overtoni or P. currii. Prognathodontins were important predators in the Maastrichtian of Africa, previously being recorded in Morocco, Congo, and Angola.

1. Introduction

The mosasaurs (Mosasauridae) are an extinct group of marine reptiles that became abundant and diverse in the Upper Cretaceous [1]. Early work on mosasaurs focused heavily on Europe and North America [2,3,4,5,6,7], where paleontology first began as a science. Later on, they were also documented in South America and Antarctica [8,9,10,11,12,13,14,15,16,17,18]. However, mosasaurs appear to have been particularly diverse in the tropics and subtropics [2], which have until recently been relatively poorly known. Far and away the most diverse mosasaurid faunas are those of Africa [19,20], with representatives of all major groups of mosasaur having been found in Africa. These include Mosasaurinae [21,22,23,24,25,26,27,28], Halisaurinae [29,30,31,32], Tylosaurinae [33], and Plioplatecarpinae [34,35,36,37], along with Pachyvaranidae [38], which may represent a primitive, basally diverging branch of Mosasauroidea. In Egypt, mosasaurids have previously been recorded from the Upper Cretaceous deposits of the Eastern Desert [39,40,41,42,43,44,45], the Nile Valley [46], and the Western Desert [47,48,49,50] (see Appendix).
This study reports a mosasaurid collected from the Maastrichtian of the Dakhla Shale, at Dakhla Oasis (Figure 1). The teeth show features suggesting affinities with Prognathodon or an allied species of Prognathodontini. The Prognathodontini are a subgroup of Mosasaurinae that include Prognathodon and its relatives, sometimes recovered as a distinct clade [24], and sometimes as close relatives of Globidens in Globidensini. More than a dozen species of Prognathodon have been identified around the world from Africa, Asia, Europe, and North America [27,51,52,53].
Anatomical abbreviations: CL, centrum length; H, height; Hyp, hypapophysis; L, length; Prz, prezygapophysis; Poz, Postzygaphophysis; W, width; W(con), condyle width; W(cot), cotyle width; H(con), condyle height; H(cot), cotyle height; S, synapophysis.

2. Materials and Methods

The studied specimens come from the Maastrichtian of the Dakhla Shale (Figure 2), and include two nearly complete marginal tooth crowns, and three cervical, nine dorsal, and three caudal vertebrae. The fossils were found in association (Figure 3), with the material being exposed and weathered prior to collection.
The specimens described here were accessioned and housed in the New Valley Vertebrate Center (NVP), New Valley University, Kharga, under N° NVP025. Fossils were consolidated using polyvinyl acetate and prepared using brushes and air scribes. Photographs were taken using a digital camera Nikon D780 (the equipment was manufactured in Tokyo, Japan and sourced from Cairo, Egypt) and lens (Af-S Nikkor 70–200 mm 1:28E F/28L ED VR), cropped with Adobe Photoshop Version 22.4.2.

3. Stratigraphy and Age

Five formations are exposed in the Dakhla Oasis, ranging in age from Upper Cretaceous to Upper Paleocene. From the base to the top, these are the Taref Sandstone, the Quseir Variegated Shales, the Duwi Formation, the Dakhla Shale, and the Tarawan Chalk. The Taref Sandstone is predominantly formed of cross-bedded, medium- to coarse-grained sandstones. It is largely unfossiliferous. The age of the Taref Sandstone is unclear. At times it has been assigned ages ranging from Turonian [55] to Maastrichtian [56], but the ages of overlying formations constrain its age to no later than Campanian. The Quseir Variegated Shales Formation lies exposed in low buttes scattered in the plain, adjacent to the northern cliff. It is well exposed in the area of Gebel Gifata and is about 30 m thick. It is subdivided into two units, designated Units I (lower) and II (upper) by Hermina et al., (1961) [57], and Mut Member (lower) and Hindaw Member (upper) by [58]. The lower unit, made of clays, is characterized by its brick-red coloration. The upper unit consists of brown and gray gypsiferous sandy clays with a distinctive glauconitic green bed in the middle, and laminated fine sandstones at the very top. Locally, the Quseir Variegated Shales is rich in plant remains, but it is generally unfossiliferous. Recently, a titanosaurian sauropod, Mansourasaurus shahinae, was recovered from the upper member of the formation north of the road between Mut and Balat [59], along with sauropod and non-avian theropod remains [60]. The Quseir Variegated Shales has few useful index fossils and so its age is uncertain, but it is typically assigned to the Campanian [49,55,59,61,62,63]. The Duwi Formation, 10–25 m thick, comprises several phosphatic layers separated by dark beds of shale. It is the most important phosphate formation of the Upper Cretaceous of Egypt. It is largely made of polished, rounded phosphatic grains of various sizes and is particularly rich in fish teeth, including both bony fishes and elasmobranchs. The Duwi Formation is dated to the upper Campanian by macroinvertebrates, mainly ammonites [64,65] and planktic foraminifera [66]. Its contact with the overlying Dakhla Shale coincides with the Campanian–Maastrichtian boundary [66]. The Dakhla Shale consists predominantly of dark gray shales and mudstones, intercalated by thin- to thick-bedded light gray and brown sandstones. The Dakhla Shale constitutes almost the entirety of the scarp face to the north of the Dakhla Oasis, attaining nearly 200 m in thickness at Gebel Gifata. It is Maastrichtian to upper Paleocene based on planktic foraminifera and calcareous nannofossils [66,67]. The Tarawan Chalk forms the top of the plateau that bounds the Dakhla oasis to the north. It is around 20 m thick. The Tarawan Chalk is upper Paleocene, based on planktic foraminifera [67].
The fossils described here were collected from the lower part of the Dakhla Shale at Gebel Gifata, some 15 m above the base of the formation. Their low stratigraphic position suggests that they are likely lower Maastrichtian in age. Tantawy et al. [66] identified, in the lower 3 m of the Dakhla Shale, a planktic foraminiferal assemblage indicative of Zone CF8b; hence the basal Maastrichtian. An interval of about 70 m above this zone is reportedly barren of both planktic foraminifera and calcareous nannofossils [66]. Above this barren interval, Tantawy et al. (2001) [66] recorded rich microfossil assemblages indicating the calcareous nannofossil Zone CC25a and the upper part of the planktic foraminiferal Zone CF7 (upper Maastrichtian).

4. Results

4.1. Systematic Paleontology

Squamata Oppel, 1811 [68];
Mosasauroidea Gervais, 1853 [69];
Mosasauridae Gervais, 1853 [69];
Mosasaurinae Gervais, 1853 [69];
Prognathodontini Russell, 1967 [2];
Prognathodon sp. Dollo, 1889 [70];
Locality. Gebel Gifata, 12 km NW of Mut, Dakhla Oasis;
Horizon. Lower part of the Dakhla Shale, lower Maastrichtian;
Material. NVP025, two marginal teeth, three cervical, nine dorsal, three caudal vertebrae, and ribs, found in association (Figure 3).

4.2. Comparative Description

4.2.1. Dentition

The dentition of NVP 025 includes two nearly complete marginal tooth crowns. The height of the more complete crown is 35 mm (Figure 4a,b); it has an aspect ratio (height/basal width) in lateral view equal to 1.4. The teeth are large, robust, and triangular in shape in the lateral view. Overall, the teeth resemble those of other Prognathodontini in having a robust, conical crown and weak curvature. In morphology, they are intermediate between the relatively sharp teeth of P. solvayi [71], P. saturator [52], and Thalassotitan atrox [24], and the blunt teeth of P. overtoni [72] and P. currii [51].
The crowns are bucco-lingually compressed, with elliptical to oval cross sections, and are slightly lingually curved. They are inflated at the base and narrow gradually to the tip. The apex is blunt, similar to P. overtoni and P. currii. The enamel is generally smooth, although there are faint anastomosing ridges and shallow wrinkles as in P. solvayi, P. giganteus [71], P. currii [51], P. lutugini [73], T. atrox [24], Globidens spp. [23,74], and Carinodens belgicus [21]. There are very faint facets extending up to two-thirds of the height of the crown.
Carinae are prominent on the anterior and posterior margins of the tooth, as is typical of Mosasaurinae. The anterior carina has a pronounced anterior curvature, and the posterior one is almost straight. Serrations are absent except for very faint serrations on the posterior carina. The carinae divide the teeth into two asymmetrical surfaces. The two surfaces are convex, with the buccal surface being wider and more convex than the lingual surfaces.

4.2.2. Postcranial Skeleton

The postcranial elements of NVP025 are represented by three cervical vertebrae, some ribs, nine dorsal vertebrae and three caudal vertebrae, found in association.
Cervical vertebrae. The smallest cervical vertebra (Figure 5) is missing its posterior end. Its centrum in the anterior view is oval-shaped, with a shallow cotyle. The prezygapophyses are large. The length of the centrum cannot be measured because it is broken; the synapophyses are also broken. Part of a v-shaped hypapophyseal peduncle is present. The neural arch has a sub-triangular shape. The neural canal, seen from the front, is oval in outline, and wider than high.
The other cervical vertebrae can be generally described as follows. The centrum is longer than it is high or wide. It is waisted, with a constriction between condyle and cotyle. The condyle and cotyle surfaces are oval in cross section, with a dorsal flat surface; they are wider than they are tall (Table 1). The condyle has a hemispherical apex, while the cotyle has shallow to moderately deep depression. The neural arch is located on the midline of the centrum and very close to the condyle face. It is dorsoventrally tall and antero-posteriorly short.
Prezygapophyses are large and oval in shape, mediolaterally compressed, and dorsoventrally steeply inclined. The prezygapophyses are approximately as tall as the anterior articular face of the vertebra. They are separated from the neural arch by a slightly long pedicle and more projected past the cotyle epiphysis. The postzygapophyses are smaller than the pre-zygapophyses and steeply inclined mediolaterally, forming elliptical shapes.
Anterior and posterior zygapophyses are well-developed. Zygosphenes and zygantra are absent as in P. overtoni [71], but unlike in P. solvayi [71], P. giganteus [71], P. saturator [52], P. lutugini [73], and Thalassotitan atrox [24], where cervical vertebrae have functional zygosphenes and zygantra.
Each cervical vertebra bears a robust, posteriorly directed hypaphophysis on the posterior half of the centrum’s ventral surface. It is long with an oval and concave facet on the ventral side (Figure 5). The synapophysis is well developed, dorsoventrally compressed, and extends posterolaterally from the centra, with the long axis forming an angle approximately 90° from the sagittal plane. The synapophysis is anteroposteriorly positioned nearly above the midline of the centra. It is oval in cross section. It is thick and wide at the base and tapers toward their lateral ends.
Dorsal vertebrae. Dorsal vertebrae (Figure 6) are distinguished from cervical vertebrae by the lack of the hypapophyseal peduncle. The centra are longer than they are tall or wide; generally, they possess sub−triangular to oval or kidney-shaped condylar articulations, which are dorsoventerally compressed (condyles are not higher than wide; see Table 2). One dorsal vertebra preserves the prezygapophyses. They are large, but smaller than those of the cervical vertebrae. The synapophyses are robust, and wider than in the cervical vertebrae. They are dorsoventrally compressed, horizontally oriented, projecting antero-posteriorly from the midline portion of the centra, and taper at their ends. The centra of the dorsal vertebrae vary in shape from oval to subtriangular, and the cotyles vary in depth from shallower to deeper, indicating that they come from different places in the body. The neural canal is smaller than in cervical vertebrae, forming a sub-triangular opening that is taller than it is wide. The dorsal spines are poorly preserved.
Caudal vertebrae. Three caudal vertebrae were found. One vertebra is well-preserved (Figure 7). They generally have long neural spines, long and broad transverse processes and lack haemal arches (have straight ventral surfaces), which may refer to the caudal vertebrae. The centra are long, unlike P. solvayi [71]. The neural arch is triangular. The condylar articulation is shallower than in dorsal vertebrae and its shape becomes more subtriangular or subcircular. The condyles and cotyles are not dorsoventrally compressed. The ratio between the height and width of the vertebral centra is 1:1. The neural arch is longer than in cervical vertebrae.

5. Discussion

5.1. Comparison

NVP025 shows features of Prognathodontini, such as the large, robust tooth crowns and the presence of smooth enamel, supporting the attribution of NVP025 to Prognathodon [75,76,77]. The teeth of NVP025 are also constricted at the base and show slight inflation just above the base that is characteristic of Prognathodon [3,75,78].
NVP025 can be distinguished from the type species of P. solvayi [70,71] in that P. solvayi teeth have vertical striae, well-defined facets [77], and more sharply pointed apices. Although the teeth of NVP025 resemble the teeth of P. overtoni [53], they differ from P. overtoni in that P. overtoni teeth have completely smooth enamel [72]. Some specimens referred to as P. overtoni have relatively sharp tooth apices [79] (Figure 4), but others have blunter teeth, more like those of NVP025 [79] (Figure 4); the significance of this variation (i.e., whether it is individual variation, ontogenetic, or interspecific) remains unclear. NVP025 differs from P. currii [20,51] (Lower Maastrichtian of Ganntour Basin, Morocco, Maastrichtian of Negev-Israel), in that P. currii teeth have blunter apices and straighter crowns, and have blunt massive carinae, unlike the sharp carinae of NVP025. P. currii also has highly rugose enamel at the tooth apex, and is a much larger animal. NVP025 differs from P. saturator [52], in that P. saturator teeth are rounded in cross-section and more inflated, and have no medial curvature; they are also generally smooth, and have blunter apices. NVP025 differs from P. giganteus [80], in that P. giganteus (Maastrichtian of Belgium) teeth are more inflated with a subcircular cross section [71]. It differs from Prognathodon lutugini [73,81] in that P. lutugini teeth have slender shaped crowns and strongly serrated carinae. It also differs from Prognathodon kianda [27], in that P. kianda teeth have completely smooth enamel and more pointed crowns. It also differs from Prognathodon waiparaensis [82] (Maastrichtian of New Zealand), in that P. waiparaensis teeth have a curved, well-serrated posterior carina and have defined facets. It differs from Prognathodon hudae [83] in that the apex of NVP025 is relatively more pointed, the teeth are more laterally compressed, and the anterior and posterior carinae are strong and pronounced in contrast with P. hudae, which has faint anterior carinae. It differs from Prognathodon compressidens [84] (Campanian of France) in that P. compressidens teeth are much smaller and more slender [27]. It differs from Prognathodon sectorius [85] in that P. sectorius teeth have straight anterior and posterior carinae with more flat lateral surface and equal labial and lingual surfaces [86,87]. It differs in shape from T. atrox [24] in that the teeth of T. atrox have a conical shape and sharp apices; meanwhile, NVP025 has more compressed teeth and blunter apices. It also differs from all Mosasaurus spp., which have well-defined faceted crowns and pointed apices. It differs from Eremiasaurus heterodontus [25], in that E. heterodontus teeth have pointed apices and well-defined facets.
NVP025 can also be distinguished by vertebral morphology when compared with other Prognathodon species, particularly for the cervicals. The cervical vertebrae show several distinctive characters, including the unusually large and steeply inclined prezygapophyses, the very large hypapohyses, and the dorso-ventrally compressed synapophyses. In the synapophyses, NVP025 differs from P. solvayi, P. gigantus, and P. lutugini [71,73], which have cervical vertebrae with synapophyses originating from the anterior-most part of the centra, while NVP025 has synapophyses that extend from the central part of the vertebra. It also differs from P. solvayi, which has a broad and very long transverse process and a relatively small neural spine of the caudal vertebrae [71].
Nearly all the vertebral processes in Mosasaurus spp. cervicals are heavily “buttressed” [88], or have wide, sloping bases, but the synapophyses in NVP025 are more “discrete” in that they arise more abruptly from the sides of the centrum. NVP025 differs from Mosasaurus hoffmannii [89] in that the space between the prezygapophyses is wider in M. hoffmannii than in NVP025 (as in the space across the midline between the prezygapophyses of a single vertebra) [88]. The prezygapophyses and hypophyses are relatively taller in NVP025 than in M. hoffmannii. Also, the transverse processes are more dorso-ventrally compressed in NVP025 than is typical of Mosasaurus spp. NVP025 cervical vertebrae differ from the cervical vertebrae of T. atrox in that the hypapophyseal peduncle is much larger in NVP025.
Overall, the morphology of the teeth and vertebrae, and the size of the animal (Figure 8), seem most similar to P. overtoni. However, because of the lack of diagnostic cranial elements, it is provisionally placed in Prognathodontini as Prognathodon sp. Future finds will hopefully provide diagnostic material for this species, either allowing a confident referral to P. overtoni or the recognition of a new species.

5.2. Implications for Prognathodon Biodiversity and Biogeography

Discoveries over the past half century have shown the widespread presence of Prognathodon in the subtropics of Africa and the Middle East, with Prognathodon now known in Egypt, Morocco, Israel, Congo, and Angola [6,24,29,90]. The discovery of the present Prognathodon specimens from Egypt increases our knowledge of the diversity and distribution of the Maastrichtian mosasaurs in Egypt and North Africa, and emphasizes the endemism of mosasaur faunas (Figure 9).

Author Contributions

Conceptualization, G.A.A.E.-K., A.A.S., H.P.S., N.R.L. and A.S.; investigation, G.A.A.E.-K., A.A.S., N.R.L. and A.S.; writing—original draft preparation, G.A.A.E.-K., A.A.S., H.P.S. and A.S.; writing—review and editing, A.A. and M.A. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by New Valley University.

Data Availability Statement

All the data used in the study are included in the paper.

Acknowledgments

The authors would like to thank the President of the New Valley University, Abdulaziz Tantawi, for his support in the field trips. We would also like to thank Mohamed Kamel, Essam Rhema, and Heba Farahat, for their efforts in the collection and restoration of the studied specimens. Many thanks to Ahmed Ibraheem for his help in editing the photos. Thanks to the anonymous reviewers for their constructive comments. We also thank Youssef El-Hadad and Ayman Mohamadeen for their hospitality and help during the field camping.

Conflicts of Interest

The authors declare no conflict of interest.

Appendix A

Table A1. Table showing previously reported Mosasauridae specimens from the Upper Cretaceous of Egypt.
Table A1. Table showing previously reported Mosasauridae specimens from the Upper Cretaceous of Egypt.
TaxonSynonymyLocationStage References
Mosasauridae indet. Eastern DesertWadi QenaTuronian?[40,42]
Mosasauridae indet. Wadi GedamiSantonian[39,40]
Halisaurus arambourgi Wadi Um HemaietCampanian[6,41]
cf. Thalassotitan atroxPrognathodon sp. novGebel DuwiCampanian[6,24,43]
Igdamanosaurus aegyptiacusGlobidens aegyptiacusUm el Huetat phosphate mines near SafagaMaastrichtian[44,94]
Prognathodon giganteus Um el Huetat phosphate mines near SafagaMaastrichtian[44,94]
Globidens phosphaticus phosphates of QuseirMaastrichtian[6,45]
Mosasauridae indet. Nile Valleyphosphate deposits of SibaiyaCampanian-Maastrichtian[6]
Mosasauridae indet. Western DesertQasr ElDakhlaMaastrichtian[47]
cf. Platecarpus Nubian Sandstone of MahamidCampanian[48]
Globidens sp. Duwi Formation near the village of Teneida, Dakhla OasisCampanian[49]
Globidens Dakhla Shale, Dakhla OasisMaastrichtian[50]
Prognathodon? sp. Dakhla Shale, Dakhla Oasis, near AL-Rashda villageMaastrichtianThis paper

Appendix B

Table A2. Comparing the characteristics of the teeth of Prognathodontini species.
Table A2. Comparing the characteristics of the teeth of Prognathodontini species.
Species CarinaeApexCross Section OrnamentationFacesShape
Character
PrognathodonsolvayiBicarinatePointed?RoundedDeeply striated Faceted or prismaticSlender
PrognathodonovertoniWeakly bicarinateBluntRoundedSmoothUnfacetedSlender, inflated
PrognathodoncurriiBicarinate, and some bear rough serrationsBluntEllipsoidSmooth enamel, fine anastomosing lines towards the apices UnfacetedSubconical, robust, straight
PrognathodonsaturatorBicarinate?BluntRoundedSmoothUnfacetedMassive, rounded
PrognathodongiganteusBicarinateBluntRoundedSmoothUnfacetedConical
PrognathodonlutuginiStrongly bicarinate with a weak serration on both carinaeBlunt?RoundedMinor wrinkles at the tip of the crownsUnfacetedBarrel-shaped
PrognathodonkiandaBicarinate, unserratedPointedRounded?Smooth, except very fine anastomosing ridgesUnfacetedSlender
PrognathodonwaiparaensisBicarinate with serration on both carinaeBluntRoundedSmoothUnfacetedConical shape
PrognathodonhudaeWeakly bicarinate and unserratedBluntSubovalSmooth with very faint anastomosing ridgesUnfacetedSlender
PrognathodonsectoriusBicarinate BluntEllipsoidSmooth and silkyUnfaceted?Slender
Thalassotitan atroxBicarinate Pointed Ornamented with coarse bumps and anastomosing ridgesUnfacetedConical in shape
Eremiasaurus heterodontusBicarinate, serrated Pointed SmoothFaint traces of facetsHigh and slender, blade-like
Prognathodon? sp.Bicarinate, unserratedBluntEllipsoidSmooth with faint anastomosing ridgesFaint traces of facetsTriangular, laterally compressed

References

  1. Polcyn, M.J.; Jacobs, L.L.; Araújo, R.; Schulp, A.S.; Mateus, O. Physical drivers of mosasaur evolution. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2014, 400, 17–27. [Google Scholar] [CrossRef]
  2. Russell, D. Systematics and morphology of American Mosasaurs (Reptilia, Sauria). Bull. Peabody Mus. Nat. Hist. 1967, 23, 1. [Google Scholar]
  3. Schulp, A.S.; Averianov, A.O.; Yarkov, A.A.; Trikolidi, F.A.; Jagt, J.W. First record of the late cretaceous durophagous mosasaur Carinodens belgicus (Squamata, Mosasauridae) from volgogradskaya oblast’(Russia) and Crimea (Ukraine). Russ. J. Herpetol. 2006, 13, 175–180. [Google Scholar]
  4. Polcyn, M.; Bell, G. Russellosaurus coheni n. gen., n. sp., a 92 million-year-old mosasaur from Texas (USA), and the definition of the parafamily Russellosaurina. Neth. J. Geosci. 2005, 84, 321–333. [Google Scholar] [CrossRef]
  5. Bardet, N.; Tunoğlu, C. The first mosasaur (Squamata) from the Late Cretaceous of Turkey. J. Vertebr. Paleontol. 2002, 22, 712–715. [Google Scholar] [CrossRef]
  6. Bardet, N. Maastrichtian marine reptiles of the Mediterranean Tethys: A palaeobiogeographical approach. Bull. Soc. Géol. Fr. 2012, 183, 573–596. [Google Scholar] [CrossRef]
  7. Bardet, N.; Corral, J.C.; Suberbiola, X.F. Les mosasaures (Squamata) du Crétacé supérieur du Bassin basco-cantabrique. Geobios 1997, 30, 19–26. [Google Scholar] [CrossRef]
  8. Fernández, M.; Martin, J.; Casadío, S. Mosasaurs (Reptilia) from the late Maastrichtian (Late Cretaceous) of northern Patagonia (Río Negro, Argentina). J. S. Am. Earth Sci. 2008, 25, 176–186. [Google Scholar] [CrossRef]
  9. Fernandez, M.; Martin, J.E. Description and phylogenetic relationships of Taniwhasaurus antarcticus (Mosasauridae, Tylosaurinae) from the upper Campanian (Cretaceous) of Antarctica. Cretac. Res. 2009, 30, 717–726. [Google Scholar] [CrossRef]
  10. Fernández, M.S.; Gasparini, Z. Campanian and Maastrichtian mosasaurs from Antarctic Peninsula and Patagonia, Argentina. Bull. Soc. Géol. Fr. 2012, 183, 93–102. [Google Scholar] [CrossRef]
  11. Fernández, M.S.; Talevi, M. An halisaurine (Squamata: Mosasauridae) from the Late Cretaceous of Patagonia, with a preserved tympanic disc: Insights into the mosasaur middle ear. Comptes Rendus Palevol 2015, 14, 483–493. [Google Scholar] [CrossRef]
  12. Gasparini, Z.; Casadío, S.; Fernández, M.; Salgado, L. Marine reptiles from the Late Cretaceous of northern Patagonia. J. S. Am. Earth Sci. 2001, 14, 51–60. [Google Scholar] [CrossRef]
  13. Martin, J.E.; Fernández, M. The synonymy of the Late Cretaceous mosasaur (Squamata) genus Lakumasaurus from Antarctica with Taniwhasaurus from New Zealand and its bearing upon faunal similarity within the Weddellian Province. Geol. J. 2007, 42, 203–211. [Google Scholar] [CrossRef]
  14. Ruiz, P.G.; Fernández, M.S.; Talevi, M.; Leardi, J.M.; Reguero, M.A. A new Plotosaurini mosasaur skull from the upper Maastrichtian of Antarctica. Plotosaurini paleogeographic occurrences. Cretac. Res. 2019, 103, 104166. [Google Scholar] [CrossRef]
  15. Salgado, L.; Fernández, M.; Talevi, M. Observaciones histológicas en reptiles marinos (Elasmosauridae y Mosasauridae) del Cretácico Tardío de Patagonia y Antártida. Ameghiniana 2007, 44, 513–523. [Google Scholar]
  16. Talevi, M.; Fernández, M.S.; Cerda, I. Osteohistología en Mosasaurios (Squamata: Mosasauridae) del Cretácico Tardío de la Cuenca James Ross (Península Antártica). Ameghiniana 2011, 48, 668–673. [Google Scholar] [CrossRef]
  17. Talevi, M.; Rothschild, B.; Fernández, M.; Reguero, M.; Mitidieri, M. A pathological scapula in a mosasaur from the upper Maastrichtian of Antarctica: Evidence of infectious arthritis and spondyloarthropathy. Cretac. Res. 2019, 100, 1–4. [Google Scholar] [CrossRef]
  18. Talevi, M.; Brezina, S. Bioerosion structures in a Late Cretaceous mosasaur from Antarctica. Facies 2019, 65, 5. [Google Scholar] [CrossRef]
  19. Mateus, O.; Polcyn, M.J.; Jacobs, L.L.; Araújo, R.; Schulp, A.S.; Marinheiro, J.; Pereira, B.; Vineyard, D. Cretaceous amniotes from Angola: Dinosaurs, pterosaurs, mosasaurs, plesiosaurs, and turtles. V J. Int. Sobre Paleontol. Dinosaur. Entorno 2012, 71–105. [Google Scholar]
  20. Bardet, N.; Houssaye, A.; Vincent, P.; Suberbiola, X.P.; Amaghzaz, M.; Jourani, E.; Meslouh, S. Mosasaurids (Squamata) from the Maastrichtian phosphates of Morocco: Biodiversity, palaeobiogeography and palaeoecology based on tooth morphoguilds. Gondwana Res. 2015, 27, 1068–1078. [Google Scholar] [CrossRef]
  21. Schulp, A.; Bardet, N.; Bouya, B. A new species of the durophagous mosasaur Carinodens (Squamata, Mosasauridae) and additional material of Carinodens belgicus from the Maastrichtian phosphates of Morocco. Neth. J. Geosci. 2009, 88, 161–167. [Google Scholar] [CrossRef]
  22. Longrich, N.R.; Bardet, N.; Schulp, A.S.; Jalil, N.-E. Xenodens calminechari gen. et sp. nov., a bizarre mosasaurid (Mosasauridae, Squamata) with shark-like cutting teeth from the upper Maastrichtian of Morocco, North Africa. Cretac. Res. 2021, 123, 104764. [Google Scholar] [CrossRef]
  23. Bardet, N.; Suberbiola, X.P.; Iarochène, M.; Amalik, M.; Bouya, B. Durophagous Mosasauridae (Squamata) from the Upper Cretaceous phosphates of Morocco, with description of a new species of Globidens. Neth. J. Geosci. 2005, 84, 167–175. [Google Scholar] [CrossRef]
  24. Longrich, N.R.; Jalil, N.-E.; Khaldoune, F.; Yazami, O.K.; Pereda-Suberbiola, X.; Bardet, N. Thalassotitan atrox, a giant predatory mosasaurid (Squamata) from the upper Maastrichtian phosphates of Morocco. Cretac. Res. 2022, 140, 105315. [Google Scholar] [CrossRef]
  25. LeBlanc, A.; Caldwell, M.; Bardet, N. A new mosasaurine from the Maastrichtian (U. cretaceous) phosphates of Morocco and the implications for the systematics of the Mosasaurinae. J. Vertebr. Paleontol. 2012, 32, 1–23. [Google Scholar] [CrossRef]
  26. Bardet, N.; Suberbiola, X.P.; Iarochene, M.; Bouyahyaoui, F.; Bouya, B.; Amaghzaz, M. Mosasaurus beaugei Arambourg, 1952 (Squamata, Mosasauridae) from the late Cretaceous phosphates of Morocco. Geobios 2004, 37, 315–324. [Google Scholar] [CrossRef]
  27. Schulp, A.S.; Polcyn, M.J.; Mateus, O.; Jacobs, L.L.; Morais, M.L. A new species of Prognathodon (Squamata, Mosasauridae) from the Maastrichtian of Angola, and the affinities of the mosasaur genus Liodon. In Proceedings of the Second Mosasaur Meeting, Hays, KS, USA, 3–6 May 2007; Fort Hays State University: Hays, KS, USA, 2008; pp. 1–12. [Google Scholar]
  28. Polcyn, M.J.; Jacobs, L.L.; Schulp, A.S.; Mateus, O. The North African Mosasaur Globidens phosphaticus from the maastrichtian of angola. Hist. Biol. 2010, 22, 175–185. [Google Scholar] [CrossRef]
  29. Longrich, N.R.; Bardet, N.; Khaldoune, F.; Yazami, O.K.; Jalil, N.-E. Pluridens serpentis, a new mosasaurid (Mosasauridae: Halisaurinae) from the Maastrichtian of Morocco and implications for mosasaur diversity. Cretac. Res. 2021, 126, 104882. [Google Scholar] [CrossRef]
  30. Bardet, N.; Pereda Suberbiola, X.; Iarochene, M.; Bouya, B.; Amaghzaz, M. A new species of Halisaurus. from the Late Cretaceous phosphates of Morocco, and the phylogenetical relationships of the Halisaurinae (Squamata: Mosasauridae). Zool. J. Linn. Soc. 2005, 143, 447–472. [Google Scholar] [CrossRef]
  31. Longrich, N.R. A new species of Pluridens (Mosasauridae: Halisaurinae) from the upper Campanian of Southern Nigeria. Cretac. Res. 2016, 64, 36–44. [Google Scholar] [CrossRef]
  32. Lingham-Soliar, T. A new mosasaur Pluridens walkeri from the Upper Cretaceous, Maastrichtian of the Iullemmeden Basin, southwest Niger. J. Vertebr. Paleontol. 1998, 18, 709–717. [Google Scholar] [CrossRef]
  33. Rempert, T.H.; Martens, B.P.; Melchers, A.P.V. First Record of a Tylosaurine Mosasaur from the Latest Cretaceous Phosphates of Morocco. Open J. Geol. 2022, 12, 883–906. [Google Scholar] [CrossRef]
  34. Arambourg, C. Les vertébrés fossiles des gisements de phosphates (Maroc-Algérie-Tunisie). Notes Mémoires Serv. Géol. Maroc 1952, 92, 1–372. [Google Scholar]
  35. Azzaroli, A.; De Giuli, C.; Torre, G.F.D. An aberrant Mosasaur from the Upper Cretaceous of North-Western Nigeria. Preliminary report. Atti Della Accad. Naz. Lincei. Cl. Sci. Fis. Mat. Natur. Rend. 1972, 52, 398–402. [Google Scholar]
  36. Lingham-Soliar, T. The mosasaur Plioplatecarpus (Reptilia, Mosasauridae) from the Upper Cretaceous of Europe. Bull. Inst. R. Sci. Nat. Belgique Sci. Terre 1994, 64, 177–211. [Google Scholar]
  37. Strong, C.R.; Caldwell, M.W.; Konishi, T.; Palci, A. A new species of longirostrine plioplatecarpine mosasaur (Squamata: Mosasauridae) from the Late Cretaceous of Morocco, with a re-evaluation of the problematic taxon ‘Platecarpus’ ptychodon. J. Syst. Palaeontol. 2020, 18, 1769–1804. [Google Scholar] [CrossRef]
  38. Houssaye, A.; Bardet, N.; Rage, J.-C.; Pereda-Suberbiola, X.; Bouya, B.; Amaghzaz, M.; Amalik, M. A review of Pachyvaranus crassispondylus Arambourg, 1952, a pachyostotic marine squamate from the latest Cretaceous phosphates of Morocco and Syria. Geol. Mag. 2011, 148, 237–249. [Google Scholar] [CrossRef]
  39. Barron, T.; Hume, W.F. Notes sur la Géologie du Désert Oriental de L’Égypte; Éditeur Inconnu: Hong Kong, 1901. [Google Scholar]
  40. Barron, T.; Hume, W.F. Topography and Geology of the Eastern Desert of Egypt, Central Portion; National Print Department: Hong Kong, 1902. [Google Scholar]
  41. Priem, F. Sur des vertébrés du Crétacé et de l’Eocène d’Egypte. Bull. L’inst. D’égypte 1914, 5, 127–128. [Google Scholar] [CrossRef]
  42. Blanckenhorn, M. Aegypten, Handbuch der regionalen Geologie; Heidelberg Carl Winters University: Heidelberg, Germany, 1921. [Google Scholar]
  43. Gemmellaro, M. Ittiodontoliti maestrichtiani di Egitto. Atti Della Accad. Sci. Lett. Arti Palermo 1920, 7, 149–204. [Google Scholar]
  44. Zdansky, O. The occurrence of mosasaurs in Egypt and in Africa in general. Bull. L’inst. D’egypte 1934, 17, 83–94. [Google Scholar] [CrossRef]
  45. Leonardi, P.; Malaroda, R. Prima segnalazione di un mosasauro del genere Globidens nel Cretaceo del’Egytto. Acta Pontif. Acad. Sci. 1946, 10, 183–188. [Google Scholar]
  46. Bardet, N. The mosasaur collections of the Muséum National d’Histoire Naturelle of Paris. Bull. Soc. Géol. Fr. 2012, 183, 35–53. [Google Scholar] [CrossRef]
  47. Quaas, A. Beitrage zur Kenntniss der Fauna der Obersten Kreidebildungen in der libyschen Wüste (Overwegischichten und Blätterthone); E. Schweizerbart: Stuttgart, Germany, 1902. [Google Scholar]
  48. Stromer, E.; Weiler, W. Beschreibung von Wirbeltier-Resten aus dem nubischen Sandsteine Oberägyptens und aus ägyptischen Phosphaten nebst Bemerkungen über die Geologie der Umgegend von Mahamîd in Oberäg. In Beschreibung von Wirbeltier-Resten aus dem Nubischen Sandsteine Oberägyptens und aus Agyptischen Phosphaten Nebst Bemerkungen uber Die Geologie der Umgegend Von Mahamîd in Oberäg ; Oldenbourg Wissenschaftsverlag: Minhen, Germany, 1930. [Google Scholar]
  49. Sallam, H.M.; O’Connor, P.M.; Kora, M.; Sertich, J.J.; Seiffert, E.R.; Faris, M.; Ouda, K.; El-Dawoudi, I.; Saber, S.; El-Sayed, S. Vertebrate paleontological exploration of the Upper Cretaceous succession in the Dakhla and Kharga Oases, Western Desert, Egypt. J. Afr. Earth Sci. 2016, 117, 223–234. [Google Scholar] [CrossRef]
  50. AbdelGawad, M.K.; El-Kheir, G.A.A.; Kassab, W.G. The youngest records of mosasaurid reptiles from the Upper Cretaceous of the South-Western Desert in Egypt. Proc. Geol. Assoc. 2021, 132, 556–562. [Google Scholar] [CrossRef]
  51. Christiansen, P.; Bonde, N. A new species of gigantic mosasaur from the Late Cretaceous of Israel. J. Vertebr. Paleontol. 2002, 22, 629–644. [Google Scholar] [CrossRef]
  52. Dortangs, R.W.; Schulp, A.S.; Mulder, E.W.; Jagt, J.W.; Peeters, H.H.; De Graaf, D.T. A large new mosasaur from the Upper Cretaceous of The Netherlands. Neth. J. Geosci. 2002, 81, 1–8. [Google Scholar] [CrossRef]
  53. Williston, S.W. Brachysaurus, a new genus of mosasaurs. Kans. Univ. Q. 1897, 6, 95–98. [Google Scholar]
  54. Abu El-Kheir, G.-M.; Abdelgawad, M.K.; Kassab, W.G. First known gigantic sea turtle from the Maastrichtian deposits in Egypt. Acta Palaeontol. Pol. 2021, 66, 349–355. [Google Scholar] [CrossRef]
  55. Hermina, M. The surroundings of Kharga, Dakhla and Farafra oases. In the Geology of Egypt; Routledge: Abingdon, UK, 2017; pp. 259–292. [Google Scholar]
  56. Said, R. The Geology of Egypt; Elsevier: Amsterdam, The Netherlands, 1962; Volume 377. [Google Scholar]
  57. Hermina, M.; Ghorbrial, M.; Issawi, B. The Geology of the Dakhla Area: Geol; Survey and Mineral Resources Department: Cairo, Egypt, 1961; pp. 1–33. [Google Scholar]
  58. Omara, S.; Philobbos, E.; Mansour, H. Contribution to the geology of the Dakhla Oasis area, Western Desert, Egypt. Bull. Fac. Sci. Assiut Univ. 1976, 5, 319–339. [Google Scholar]
  59. Sallam, H.M.; Gorscak, E.; O’Connor, P.M.; El-Dawoudi, I.A.; El-Sayed, S.; Saber, S.; Kora, M.A.; Sertich, J.J.; Seiffert, E.R.; Lamanna, M.C. New Egyptian sauropod reveals Late Cretaceous dinosaur dispersal between Europe and Africa. Nat. Ecol. Evol. 2018, 2, 445–451. [Google Scholar] [CrossRef]
  60. Salem, B.S.; O’Connor, P.M.; Gorscak, E.; El-Sayed, S.; Sertich, J.J.; Seiffert, E.; Sallam, H.M. Dinosaur remains from the Upper Cretaceous (Campanian) of the Western Desert, Egypt. Cretac. Res. 2021, 123, 104783. [Google Scholar] [CrossRef]
  61. Barthel, K.W. Late Cretaceous and Early Tertiary Stratigraphy in the Great Sand Sea and Its SE Margins (Farafra and Dakhla Oases), SW Desert, Egypt; Pascual Francis: Lawrence, KS, USA, 1981. [Google Scholar]
  62. Luger, P.; Schrank, E. Mesozoic to Paleogene transgressions in middle and southern Egypt. Summary of paleontological evidence. In Colloquium on African Geology; Pascual Francis: Lawrence, KS, USA, 1987; Volume 14, pp. 199–202. [Google Scholar]
  63. Zaghloul, E.A. Geology of Dakhla Oasis, Western Desert, Egypt. In Sustainable Water Solutions in the Western Desert, Egypt: Dakhla Oasis; Springer: Berlin/Heidelberg, Germany, 2021; pp. 29–44. [Google Scholar]
  64. El Naggar, Z. Stratigraphy and planktonic foraminifera in the Upper Cretaceous–Lower Tertiary succession in the Esna-Idfu region. Nile Val. Egypt UAR Br. Mus. Nat. Hist. Bull. 1966, 2, 291. [Google Scholar]
  65. Dominik, W.; Schaal, S. Notes on the Stratigraphy of the Upper Cretaceous Phosphates (Campanian) of the Western Desert (Egypt); Pascual Francis: Lawrence, KS, USA, 1984. [Google Scholar]
  66. Tantawy, A.; Keller, G.; Adatte, T.; Stinnesbeck, W.; Kassab, A.; Schulte, P. Maastrichtian to Paleocene depositional environment of the Dakhla Formation, Western Desert, Egypt: Sedimentology, mineralogy, and integrated micro-and macrofossil biostratigraphies. Cretac. Res. 2001, 22, 795–827. [Google Scholar] [CrossRef]
  67. Hewaidy, A.G.A.; Farouk, S.; Bazeen, Y.S. Sequence stratigraphy of the maastrichtian-paleocene succession at the dakhla oasis, western desert, Egypt. J. Afr. Earth Sci. 2017, 136, 22–43. [Google Scholar] [CrossRef]
  68. Oppel, M. Die Ordnungen, Familien und Gattungen der Reptilien Als Prodrom Einer Naturgeschichte Derselben. In Commission bey J. Lindauer: Minhen, Germany, 1811. [Google Scholar]
  69. Gervais, P. Observations relatives aux reptiles fossiles de France. Comptes Rendus Hebd. Séances L’acad. Sci. 1853, 36, 470. [Google Scholar]
  70. Dollo, L. Note sur les vertébrés récemment offerts au Musée de Bruxelles par M. Alfred Lemonnier. Bull. Soc. Belg. Géol. Paléontol. D’hydrol. 1889, 3, 181–182. [Google Scholar]
  71. Lingham-Soliar, T.; Nolf, D. The mosasaur Prognathodon (Reptilia, Mosasauridae) from the Upper Cretaceous of Belgium. In Bulletin van het Koninklijk Belgisch Instituut voor Natuurwetenschappen. Aardwetenschappen = Bulletin de l’Institut Royal des Sciences Naturelles de Belgique; Sciences de la Terre KBIN: Brussel, Belgium, 1989. [Google Scholar]
  72. Konishi, T.; Brinkman, D.; Massare, J.A.; Caldwell, M.W. New exceptional specimens of Prognathodon overtoni (Squamata, Mosasauridae) from the upper Campanian of Alberta, Canada, and the systematics and ecology of the genus. J. Vertebr. Paleontol. 2011, 31, 1026–1046. [Google Scholar] [CrossRef]
  73. Grigoriev, D. Redescription of Prognathodon lutugini (Squamata, Mosasauridae). Труды Зooлoгическoгo Института РАН 2013, 317, 246–261. [Google Scholar] [CrossRef]
  74. Gilmore, C.W. A new mosasauroid reptile from the Cretaceous of Alabama. In Proceedings of the United States National Museum; U. S. National Museeum: Washington, DC, USA, 1912; Available online: https://repository.si.edu/handle/10088/14313 (accessed on 28 July 2023).
  75. Bell Jr, G.L. A phylogenetic revision of North American and Adriatic Mosasauroidea. In Ancient Marine Reptiles; Elsevier: Amsterdam, The Netherlands, 1997; pp. 293–332. [Google Scholar]
  76. Bell, G.; Polcyn, M. Dallasaurus turneri, a new primitive mosasauroid from the Middle Turonian of Texas and comments on the phylogeny of Mosasauridae (Squamata). Neth. J. Geosci. 2005, 84, 177–194. [Google Scholar] [CrossRef]
  77. Lindgren, J.; Jagt, J. Danish mosasaurs. Neth. J. Geosci. 2005, 84, 315–320. [Google Scholar] [CrossRef]
  78. Bardet, N.; Cappetta, H.; Suberbiola, X.P.; Mouty, M.; Al Maleh, A.; Ahmad, A.; Khrata, O.; Gannoum, N. The marine vertebrate faunas from the Late Cretaceous phosphates of Syria. Geol. Mag. 2000, 137, 269–290. [Google Scholar] [CrossRef]
  79. Konishi, T.; Caldwell, M.W. Two new plioplatecarpine (Squamata, Mosasauridae) genera from the Upper Cretaceous of North America, and a global phylogenetic analysis of plioplatecarpines. J. Vertebr. Paleontol. 2011, 31, 754–783. [Google Scholar] [CrossRef]
  80. Dollo, L. Les Mosasauriens de la Belgique; Vlaams Instituut voor de Zee: Oostende, Belgium, 1904. [Google Scholar]
  81. Yakovlev, N. Remains of the Late Cretaceous mosasaur from the south of Russia. Izv. Geol. Kom. 1901, 20, 507–522. [Google Scholar]
  82. Welles, S.P. Late cretaceous marine reptiles of New Zealand. Rec. Canterb. Mus. 1971, 9, 1–111. [Google Scholar]
  83. Kaddumi, H.F. Fossils of the Harrana Fauna and the Adjacent Areas; Eternal River Museum of Natural History the Hashemite Kingdom of Jordan: Tokyo, Japan, 2009. [Google Scholar]
  84. Gaudry, A. Les Pythonomorphes de France; Baudry: Domblans, France, 1892; Volume 3. [Google Scholar]
  85. Cope, E.D. Synopsis of the Extinct Batrachia and Reptilia of North America; American Philosophical Society: Philadelphia, PA, USA, 1871; Volume 14. [Google Scholar]
  86. Bardet, N.; Suberbiola, X.P.; Corral, J.-C.; Baceta, J.I.; Torres, J.Á.; Botantz, B.; Martin, G. A skull fragment of the mosasaurid Prognathodon cf. sectorius from the Late Cretaceous of Navarre (Basque-Cantabrian Region). Bull. Soc. Géol. Fr. 2012, 183, 117–121. [Google Scholar] [CrossRef]
  87. Bardet, N.; Martin, G.; Corral, J.C.; Suberbiola, X.P.; Astibia, H. New mosasaurid teeth (Reptilia: Squamata) from the Maastrichtian of Albaina (Laño quarry, condado de Treviño). Span. J. Palaeontol. 2020, 28, 69–78. [Google Scholar] [CrossRef]
  88. Street, H.P.; Caldwell, M.W. Rediagnosis and redescription of Mosasaurus hoffmannii (Squamata: Mosasauridae) and an assessment of species assigned to the genus. Mosasaurus Geol. Mag. 2017, 154, 521–557. [Google Scholar] [CrossRef]
  89. Mantell, G.X. A Tabular Arrangement of the Organic Remains of the County of Sussex. Trans. Geol. Soc. Lond. 1829, 3, 201–216. [Google Scholar] [CrossRef]
  90. Longrich, N.R.; Jalil, N.-E.; Pereda-Suberbiola, X.; Bardet, N. Stelladens mysteriosus: A Strange New Mosasaurid (Squamata) from the Maastrichtian (Late Cretaceous) of Morocco. Fossils 2023, 1, 2–14. [Google Scholar] [CrossRef]
  91. Cappetta, H.; Bardet, N.; Suberbiola, X.P.; Adnet, S.; Akkrim, D.; Amalik, M.; Benabdallah, A. Marine vertebrate faunas from the Maastrichtian phosphates of Benguérir (Ganntour Basin, Morocco): Biostratigraphy, palaeobiogeography and palaeoecology. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2014, 409, 217–238. [Google Scholar] [CrossRef]
  92. Lingham-Soliar, T. First record of mosasaurs from the Maastrichtian (Upper Cretaceous) of Zaire. PalZ 1994, 68, 259–265. [Google Scholar] [CrossRef]
  93. Schulp, A.; Polcyn, M.; Mateus, O.; Jacobs, L. Two rare mosasaurs from the Maastrichtian of Angola and the Netherlands. Neth. J. Geosci. 2013, 92, 3–10. [Google Scholar] [CrossRef]
  94. Lingham-Soliar, T. Mosasaurs from the upper Cretaceous of Niger. Palaeontology 1991, 34, 653–670. [Google Scholar]
Fossstud 01 00007 g001
Figure 1. Geological map of Dakhla area, showing the location of Gebel Gifata, 12 km NW of Mut, Dakhla Oasis (modified after [54]).
Figure 1. Geological map of Dakhla area, showing the location of Gebel Gifata, 12 km NW of Mut, Dakhla Oasis (modified after [54]).
Fossstud 01 00007 g001
Fossstud 01 00007 g002
Figure 2. Stratigraphic section of Gebel Gifata (modified after [54]).
Figure 2. Stratigraphic section of Gebel Gifata (modified after [54]).
Fossstud 01 00007 g002
Fossstud 01 00007 g003
Figure 3. Photo of the associated vertebrae of Prognathodon sp. (blue arrows).
Figure 3. Photo of the associated vertebrae of Prognathodon sp. (blue arrows).
Fossstud 01 00007 g003
Fossstud 01 00007 g004
Figure 4. Prognathodon sp., NVP025, Gebel Gifata, Dakhla Oasis, Western Desert (Egypt), lower Maastrichtian. Teeth in lingual (a,f), buccal (b,g), anterior (c,h), posterior (d,i), and apical (e,j) views, basal cross section (k). Figure shows tooth carina and wrinkles of the crown and shape of basal cross section. Bar scale = 2 cm.
Figure 4. Prognathodon sp., NVP025, Gebel Gifata, Dakhla Oasis, Western Desert (Egypt), lower Maastrichtian. Teeth in lingual (a,f), buccal (b,g), anterior (c,h), posterior (d,i), and apical (e,j) views, basal cross section (k). Figure shows tooth carina and wrinkles of the crown and shape of basal cross section. Bar scale = 2 cm.
Fossstud 01 00007 g004
Fossstud 01 00007 g005
Figure 5. Prognathodon sp., NVP025, Gebel Gifata, Dakhla Oasis, Western Desert (Egypt), lower Maastrichtian. Cervical vertebrae in anterior (a), posterior (c), lateral (b,d,e) views. Abbreviations: cdl, condyle; ctl, cotyle; hyp, hypapophysis; nc, neural canal; poz, postzygapophysis; prz, prezygapophysis; syn, synapapophysis. Bar scale = 2 cm.
Figure 5. Prognathodon sp., NVP025, Gebel Gifata, Dakhla Oasis, Western Desert (Egypt), lower Maastrichtian. Cervical vertebrae in anterior (a), posterior (c), lateral (b,d,e) views. Abbreviations: cdl, condyle; ctl, cotyle; hyp, hypapophysis; nc, neural canal; poz, postzygapophysis; prz, prezygapophysis; syn, synapapophysis. Bar scale = 2 cm.
Fossstud 01 00007 g005
Fossstud 01 00007 g006
Figure 6. Prognathodon sp., NVP025, Gebel Gifata, Dakhla Oasis, Western Desert (Egypt), lower Maastrichtian. Dorsal vertebrae in posterior (a), anterior (b,c), and dorsal (d) views. Abbreviations: cdl, condyle; ctl, cotyle; nc, neural canal; prz, prezygapophysis; syn, synapapophysis. Bar scale = 2 cm.
Figure 6. Prognathodon sp., NVP025, Gebel Gifata, Dakhla Oasis, Western Desert (Egypt), lower Maastrichtian. Dorsal vertebrae in posterior (a), anterior (b,c), and dorsal (d) views. Abbreviations: cdl, condyle; ctl, cotyle; nc, neural canal; prz, prezygapophysis; syn, synapapophysis. Bar scale = 2 cm.
Fossstud 01 00007 g006
Fossstud 01 00007 g007
Figure 7. Prognathodon sp., NVP025, Gebel Gifata, Dakhla Oasis, Western Desert (Egypt), lower Maastrichtian. Caudal vertebrae in anterior (a) and posterior (b) views. Abbreviations: cdl, condyle; ctl, cotyle; nc, neural canal; ns, neural spine; syn, synapapophysis. Bar scale = 2 cm.
Figure 7. Prognathodon sp., NVP025, Gebel Gifata, Dakhla Oasis, Western Desert (Egypt), lower Maastrichtian. Caudal vertebrae in anterior (a) and posterior (b) views. Abbreviations: cdl, condyle; ctl, cotyle; nc, neural canal; ns, neural spine; syn, synapapophysis. Bar scale = 2 cm.
Fossstud 01 00007 g007
Fossstud 01 00007 g008
Figure 8. Silhouette showing the approximate size of Prognathodon sp. Scale = 5 m.
Figure 8. Silhouette showing the approximate size of Prognathodon sp. Scale = 5 m.
Fossstud 01 00007 g008
Fossstud 01 00007 g009
Figure 9. Map showing distribution of prognathodontins in Africa during the Upper Cretaceous. 1, Ouled Abdoun Basin, Morocco, Thalassotitan atrox; 2, Ganntour Basin, Morocco, Thalassotitan atrox, Prognathodon currii, Prognathodon giganteus [20,91]; 3, Gebel Duwi, Egypt, cf. Thalassotitan [43]; 4, Gebel Gifata, Egypt, Prognathodon sp. (this paper); 5, Congo, Prognathodon cf. giganteus [92]; 6, Namibe basin, Angola, Prognathodon kianda, Prognathodon cf. saturator [27,93]; 7, Cabinda basin, Angola, Prognathodon sp., Prognathodon cf. currii [6]. Map redrawn after map by Ron Blakey.
Figure 9. Map showing distribution of prognathodontins in Africa during the Upper Cretaceous. 1, Ouled Abdoun Basin, Morocco, Thalassotitan atrox; 2, Ganntour Basin, Morocco, Thalassotitan atrox, Prognathodon currii, Prognathodon giganteus [20,91]; 3, Gebel Duwi, Egypt, cf. Thalassotitan [43]; 4, Gebel Gifata, Egypt, Prognathodon sp. (this paper); 5, Congo, Prognathodon cf. giganteus [92]; 6, Namibe basin, Angola, Prognathodon kianda, Prognathodon cf. saturator [27,93]; 7, Cabinda basin, Angola, Prognathodon sp., Prognathodon cf. currii [6]. Map redrawn after map by Ron Blakey.
Fossstud 01 00007 g009
Table 1. Measurements of cervical vertebrae, in millimeters (mm).
Table 1. Measurements of cervical vertebrae, in millimeters (mm).
Vertebra No.W(con)W(cot)H(con)H(cot)CL S HYPPrzPoz
HWL
1--50.38--36.43--------------
248.7449.233.3237.0874.7540.4823.9433.0526.4340.8020.91
3--51.5--39.02--43.7531.452935025.54----
Table 2. Measurements of dorsal vertebrae, in millimeters (mm).
Table 2. Measurements of dorsal vertebrae, in millimeters (mm).
Vertebra No.W(con)W(cot)H(con)H(cot)CLS
1--72.57--51.3079.9543.52
264.5565.1845.9746.8276.1344.6
3--64.79--47.12--57.38
4--59.49--41.63--50.79
567.50--51.41----47.21
6--71.92--------
7--71.11--47.81----
870.34--53.01--78.2152.59
9--58.30--38.3670.71--
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.

Share and Cite

MDPI and ACS Style

El-Kheir, G.A.A.; Shaker, A.A.; Street, H.P.; Longrich, N.R.; Strougo, A.; Asan, A.; AbdelGawad, M. A Prognathodontin Mosasaur from the Maastrichtian of the Dakhla Oasis, Western Desert, Egypt. Foss. Stud. 2023, 1, 60-75. https://doi.org/10.3390/fossils1010007

AMA Style

El-Kheir GAA, Shaker AA, Street HP, Longrich NR, Strougo A, Asan A, AbdelGawad M. A Prognathodontin Mosasaur from the Maastrichtian of the Dakhla Oasis, Western Desert, Egypt. Fossil Studies. 2023; 1(1):60-75. https://doi.org/10.3390/fossils1010007

Chicago/Turabian Style

El-Kheir, Gebely A. Abu, Ahmed A. Shaker, Hallie P. Street, Nicholas R. Longrich, Amin Strougo, Anhar Asan, and Mohamed AbdelGawad. 2023. "A Prognathodontin Mosasaur from the Maastrichtian of the Dakhla Oasis, Western Desert, Egypt" Fossil Studies 1, no. 1: 60-75. https://doi.org/10.3390/fossils1010007

APA Style

El-Kheir, G. A. A., Shaker, A. A., Street, H. P., Longrich, N. R., Strougo, A., Asan, A., & AbdelGawad, M. (2023). A Prognathodontin Mosasaur from the Maastrichtian of the Dakhla Oasis, Western Desert, Egypt. Fossil Studies, 1(1), 60-75. https://doi.org/10.3390/fossils1010007

Article Metrics

Back to TopTop