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. 2020 Apr 21:928:1-216.
doi: 10.3897/zookeys.928.47517. eCollection 2020.

Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco

Nizar Ibrahim  1 , Paul C Sereno  2 , David J Varricchio  3 , David M Martill  4 , Didier B Dutheil  5 , David M Unwin  6 , Lahssen Baidder  7 , Hans C E Larsson  8 , Samir Zouhri  9 , Abdelhadi Kaoukaya  7
Affiliations

Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco

Nizar Ibrahim et al. Zookeys. .

Abstract

The geological and paleoenvironmental setting and the vertebrate taxonomy of the fossiliferous, Cenomanian-age deltaic sediments in eastern Morocco, generally referred to as the "Kem Kem beds", are reviewed. These strata are recognized here as the Kem Kem Group, which is composed of the lower Gara Sbaa and upper Douira formations. Both formations have yielded a similar fossil vertebrate assemblage of predominantly isolated elements pertaining to cartilaginous and bony fishes, turtles, crocodyliforms, pterosaurs, and dinosaurs, as well as invertebrate, plant, and trace fossils. These fossils, now in collections around the world, are reviewed and tabulated. The Kem Kem vertebrate fauna is biased toward large-bodied carnivores including at least four large-bodied non-avian theropods (an abelisaurid, Spinosaurus, Carcharodontosaurus, and Deltadromeus), several large-bodied pterosaurs, and several large crocodyliforms. No comparable modern terrestrial ecosystem exists with similar bias toward large-bodied carnivores. The Kem Kem vertebrate assemblage, currently the best documented association just prior to the onset of the Cenomanian-Turonian marine transgression, captures the taxonomic diversity of a widespread northern African fauna better than any other contemporary assemblage from elsewhere in Africa.

Keywords: Africa; Cretaceous; Douira Formation; Gara Sbaa Formation; dinosaur; paleoenvironment; vertebrate.

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Figures

Figure 1.
Figure 1.
Geographical setting of the Kem Kem region and outcrops. A View of the position of Morocco in Africa and location of the Kem Kem beds (shown in red). B Map showing the geographical location of the Kem Kem in North Africa relative to roughly coeval sites in northern Africa. C Cretaceous outcrops along the Kem Kem and Guir Hamadas (modified from Sereno et al. 1996). Numbers: 1 Kem Kem, Morocco. 2 Gara Samani, Algeria. 3 Timimoun, Algeria. 4 Monts des Ksours, Algeria. 5 Djoua Valley, Algeria. 6 Al Hamra Hamada, Libya. 7 In Abangharit, Niger. 8 Bahariya, Egypt, 9 Tataouine, Tunisia.
Figure 2.
Figure 2.
Geographical setting of the Kem Kem region. Cenomanian (~94 Mya) paleogeographic world map showing key localities (map after Scotese 2002, da Silva and Gallo 2007). Abbreviations: 1 Kem Kem region in Africa 2 Tethys Ocean 3 opening Atlantic Ocean 4 South America.
Figure 3.
Figure 3.
Outcrops of the Kem Kem sequence near Gara Sbaa. The red beds are overlain by a Cenomanian-Turonian limestone platform.
Figure 4.
Figure 4.
Example Kem Kem localities. A Basal outcrop at Aferdou N’Chaft B Iferda N’Ahouar C Gara Sbaa D Other outcrops at Gara Sbaa E Outcrops south of Jbel Zireg F Outcrop at Moher (south of Tafraoute) near the Morocco-Algeria border.
Figure 5.
Figure 5.
Outcrop near Boumerade, identified in 2008. A Flat erosional surface yielding fossils (turtle and pterosaur) B Sloped erosional surfaces yielding fossils (turtle). Abbreviations: 1 Collecting surface 2 Boundary between upper and lower members 3 Locality of a partial turtle carapace.
Figure 6.
Figure 6.
Examples of specimen measurements. A Turtle carapace fragment (MRS 172, MNHN specimen) B Femur (MPDM 270) in anterior and lateral view. Scale bars equal 2 cm in A and 10 cm in B. Abbreviations: 1 Length of specimen 2 Width measurement 3 Maximum length measurement 4 Depth measurement.
Figure 7.
Figure 7.
Outcrops of the "Continental intercalaire infracenomanien" in the Kem Kem (near Gara Sbaa). Lines separate the Cenomanian-Turonian platform that caps the "Continental intercalaire" from the upper and lower units.
Figure 8.
Figure 8.
Schematic and simplified geological cross section from the southern Atlas mountains at Amellagou to the northern margin of the Hamada du Kem Kem at Ouzina spanning the Tafilalt Platform. Notice that the Kem Kem Group strata overlie a thick Mesozoic sequence in the north within the southern Atlas Mountain thrust belt but rest unconformably on the Paleozoic basement in the south. Likely the Tafialt Platform was an 'island' within the gigantic Kem Kem river system. The line of sections extends from A at Amellagou to B at Ouzina.
Figure 9.
Figure 9.
Landsat image of southeastern Morocco showing key localities, villages and cities, modified from Ibrahim et al. 2010. Scale bar equals 25 km. Abbreviations: 1 Tiknioune Bou Tazoult (just east of Zguilma), 2 Talidat, 3 Jbel Sdila, 4 Iferda Timenkhirt, 5 Gara Sbaa, 6 Oum Tkout, 7 Boumerade, 8 Moher, 9 locality just south of Zireg, 10 Daoura, 11 locality of the neotype of Carcharodontosaurus saharicus (Sereno et al. 1996), 12 Iferda N’Ahouar west, 13 Iferda N’Ahouar east, 14 Aferdou N’Chaft, 15 Hmar Lakhdad, 16 Douira Tagemout.
Figure 10.
Figure 10.
Jagged limestone talus from the overlying Cenomanian-Turonian platform at Gara Sbaa obscures the majority of the outcrop of the Kem Kem Group.
Figure 11.
Figure 11.
The escarpment at Gara Sbaa serves as a geographic landmark in the Kem Kem region.
Figure 12.
Figure 12.
Paleozoic strata in the Kem Kem and Tafilalt regions. A Unconformity at Iferda N’Ahouar B Devonian outcrop at Hmar Lakhdad (Fig. 9: locality 15) C Geopetal infill structures in Devonian strata near Hmar Lakhdad. Abbreviations: K Cretaceous Pz Paleozoic.
Figure 13.
Figure 13.
Tectonic setting of the Kem Kem region. A Major tectonic complexes in southeastern Morocco (modified from Baidder 2007) B Kem Kem Hamada fault lines (top unnamed, bottom Omjrane-Taouz fault, Google Imagerie 2010 Digital Globe). Abbreviations: AAMF Anti Atlas Major Fault EF Erfoud Fault F fault line TF Taouz Fault.
Figure 14.
Figure 14.
Continental red beds at Aoufous.
Figure 15.
Figure 15.
Generalized lithostratigraphic sections of the Kem Kem Group from Douira in the north to Hassi Zguilma in the south. Vertical scale bar equals 100 m.
Figure 16.
Figure 16.
Northern lithostratigraphic sections of the Kem Kem Group. Vertical scale bar equals 100 m. Abbreviations: c clay p pebble sd sand st silt.
Figure 17.
Figure 17.
Southern lithostratigraphic sections of the Kem Kem Group. Vertical scale bar equals 100 m. Abbreviations: c clay p pebble sd sand st silt.
Figure 18.
Figure 18.
Outcrop of Paleozoic rocks underlying the Kem Kem Group near Hmar Lakhdad.
Figure 19.
Figure 19.
Paleozoic rocks underlying the Kem Kem Group. A Jbel Sdila locality. B Close-up of Paleozoic basement.
Figure 20.
Figure 20.
The Kem Kem Group at Iferda Timenkherin. A Overview B Close-up view of the boundary between members at Gara Sbaa: 1 Quaternary aeolian sand 2 Lower member 3 Boundary between the upper and lower members. Abbreviations: DF Douira Formation GSF Gara Sbaa Formation.
Figure 21.
Figure 21.
Limit (arrow) between the Gara Sbaa and Douira Formations.
Figure 22.
Figure 22.
Large-scale primary sedimentary structures. Large sedimentary structures occur occasionally in the upper parts of the Gara Sbaa Formation and the lower parts of the Douira Formation. In contrast to the primarily sandstone unit in (A), the lithology of (B) consists of inter-bedded sandstone and finer-grained beds. Human for scale in each equals 185 cm. The large-scale cross-bedding with good bottomsets and relatively low angle foresets is consistent with small Gilbert-type delta fronts. Alternatively, they could represent large point bar deposition with the differing lithologies of B as inclined heterolithic stratification.
Figure 23.
Figure 23.
Primary sedimentary structures in the Gara Sbaa Formation. A Linear cross-bedding at Gara Sbaa B Trough cross-bedding at Aferdou N’Chaft C Shallow channel cross-bedding (arrow) at Gara Sbaa D Tabular cross-bedding from unidirectional flow at Gara Sbaa E Large-scale cross-bedding at Gara Sbaa F Trough cross-bedding at Aferdou N’Chaft.
Figure 24.
Figure 24.
Tabular cross-bedding in the Gara Sbaa Formation. A Weathering of common tabular cross-beds B Close-up view of tabular cross-beds.
Figure 25.
Figure 25.
Sediment deformation and surface structures in the Gara Sbaa Formation at Gara Sbaa. A Large-scale overturned cross-bedding B Cross-section of overturned cross-bedding. C Surface view of overturned cross-bedding D Ripple marks E Desiccation cracks.
Figure 26.
Figure 26.
Enigmatic concretions (‘kerkoubs’) in the Gara Sbaa Formation. A In cross-stratified sandstones at Gara Sbaa B In flow direction near Boumerade C With linear sedimentary structures at Gara Sbaa D With great size variation at Gara Sbaa E In tabular cross-bedding at Gara Sbaa F In high density.
Figure 27.
Figure 27.
Close-up of ‘kerkoubs’ showing granular surface texture. Scale bar equals 1 cm.
Figure 28.
Figure 28.
Exposures of the Douira Formation and underlying sandstones. A Outcrop at Iferda N’Ahouar B Outcrop at Tikniouine Bou Tazoult. Abbreviations: 1 Limestone platform 2 Douira Formation 3 Sandstone of the Gara Sbaa Formation.
Figure 29.
Figure 29.
Satellite image at Gara Sbaa (Photo credit: Google Imagerie, 2010 Digital Globe). Scale bar equals 500 m. Abbreviations: 1 Kem Kem Group 2 Overlying limestone platform of the Akerbous Formation (Cenomanian-Turonian).
Figure 30.
Figure 30.
Primary sedimentary structures in the Douira Formation. A Fine-grained interbedding of the upper portion of the Douira Formation underlying the Cenomanian-Turonian limestone B Fine-grained mudrock with vertical paleosol zonation and root traces.
Figure 31.
Figure 31.
Oum Tkout locality where quarrying has recovered an abundance of exceptionally preserved aquatic vertebrates and invertebrates in a still-water setting.
Figure 32.
Figure 32.
Schematic paleoenvironmental stages depicting the Kem Kem region during the Cretaceous. Stages: 1 wide rivers 2 large river systems with substantial sandbanks 3 deltaic conditions 4 rise of the limestone platform.
Figure 33.
Figure 33.
Schematic geography of the major rivers systems in northern and western Africa during the Late Cretaceous (modified from Zellouf and Delfaud 1986, Delfaud 1986, Dutheil 2000). A Kem Kem river system with flow to the north B Kem Kem, paleo-Nile and paleo-Niger river systems. Abbreviations: 1 Meseta 2 Anti-Atlas 3 Ougarta 4 Kem Kem river and delta 5 Paleo-Nile river and delta 6 Paleo-Niger river and delta.
Figure 34.
Figure 34.
Geology and paleontology at the boundary between the Douira Formation and the Gara Sbaa Member of the Cenomanian-Turonian limestone at Gara Sbaa. A Marly limestone separating the Kem Kem Group from the Cenomanian-Turonian limestone B Close-up of a basal limestone unit in the Gara Sbaa Member C Oyster fossil in situ in the basal limestone D Deep-bodied teleost (Diplomystus sp.) from the Gara Sbaa Member of the Cenomanian-Turonian limestone E Long-bodied teleost (Agoultichthys chattertoni) showing preservation of soft fin structures from the Gara Sbaa Member (Murray and Wilson 2009) D and E from Martill et al. (2011). Scale bars equal 1 cm in D and 2 cm in E.
Figure 35.
Figure 35.
Fissile weathering of the Cenomanian-Turonian limestone platform.
Figure 36.
Figure 36.
Pebble and fossil lag deposits in the Kem Kem Group. A Common pebble lag deposit composed of two beds B Abraded vertebrate remains including a partial sawfish rostral tooth (Onchopristus numidus). Scale bars equal 60 cm (10 cm intervals) in A and 5 cm in B.
Figure 37.
Figure 37.
Taphonomy of Kem Kem vertebrate bone and teeth. A Articulated fish vertebrae in cross-section at Boumerade. B Cf. Spinosaurus tooth (arrow; FSAC-KK 201) adjacent to rounded pebbles C Concentration of broken bone fragments (<10 cm) D Partial archosaur tooth (MNHN-MRS 1280).
Figure 38.
Figure 38.
Taphonomy of Kem Kem vertebrate bone and teeth. A Isolated theropod quadrate (‘valley near Boumerade’ locality) B Fragmentary archosaur teeth (Gara Sbaa) C Cf. Spinosaurus vertebral zygapophysis (Gara Sbaa) D Abraded sawfish (Onchopristis numidus) rostral tooth E Isolated mixed sample of small (<1 cm) fossil vertebrates (Boumerade) F Lungfish toothplate (Boumerade). Abbreviations: 1 Turtle carapace fragment, 2 Fish vertebra, 3 Calcified rostral cartilage of Onchopristis numidus.
Figure 39.
Figure 39.
Surface and in situ collecting. A Surface collecting at Gara Sbaa B Collecting in situ fossils at Boumerade.
Figure 40.
Figure 40.
Taphonomy at Oum Tkout. A Indeterminate paraclupeid fish with mineralized muscles (white) B Soft tissue (muscle) preservation under high magnification C Complete decapod (UCRC PNV2). Scale bars equal 5 mm in A, 10 μm in B, 3 cm in C.
Figure 41.
Figure 41.
Invertebrate boring structures in dinosaur bone. A Large bone piece showing local concentration of burrows (MNHN-MRS 1568) B Close-up view of two burrows on large bone piece C Burrows of diverse diameter (UCRC I1). Scale bars equal 5 cm in A and 3 cm in C.
Figure 42.
Figure 42.
Sedimentary matrix associated with Kem Kem fossils. A Well-sorted, fine-grained sandstone on vertebrate bone (Gara Sbaa) B More oxidized coarse-grained matrix with the base of a rostral tooth of Onchopristis numidus (“valley near Boumerade”) C Moderately-cemented, red-colored matrix on an isolated archosaur bone (MNHN, unnumbered). Scale bars equal 3 cm in A, 2 cm in B, 5 cm in C.
Figure 43.
Figure 43.
Color and size variation in Kem Kem vertebrate fossils. A Theropod distal quadrate (likely from Douira) B Archosaur bone and scute fragments (collected by R. Lavocat, Kouah Trick) C Theropod left scapula (MPDM 40, unknown locality) D Cf. Rebbachisaurus partial dorsal vertebra (NMC 50844, unknown locality) E Small and medium-sized fossil fragments (UCRC unnumbered) F Sieved microfossils (UCRC unnumbered). Scale bars equal 5 cm in A and B, 10 cm in C, 20 cm in D, 5 cm in E, 3 cm in F.
Figure 44.
Figure 44.
Abrasion of Kem Kem fossils (FSAC-KK collection). A All localities B Gara Sbaa. X-axis is the abrasion index (0-4; see Table 3); Y-axis is the number of specimens. Specimen counts in A: 0 707 1 1207 2 647 3 439 4 195. Specimen counts in B: 0 158 1 433 2 396 3 330 4 138.
Figure 45.
Figure 45.
Abrasion of Kem Kem fossils (FSAC-KK collection). A Boumerade. B Aferdou N’Chaft. X-axis is the abrasion index (0-4; see Table 3); Y-axis is the number of specimens. Specimen counts in A: 0 415 1 652 2 178 3 81 4 44. Specimen counts in B: 0 71 1 41 2 14 3 7 4 5.
Figure 46.
Figure 46.
Abrasion of Kem Kem fossils (FSAC-KK collection). A Iferda N’Ahouar B Valley near Boumerade. X-axis is the abrasion index (0-4; see Table 3); Y-axis is the number of specimens. Specimen counts in A: 0 37, 1 58, 2 40, 3 13, 4 4. Specimen counts in B: 0 17, 1 15, 2 11, 3 4, 4 3.
Figure 47.
Figure 47.
Size of vertebrate fossil elements found in the Kem Kem Group (FSAC-KK collection). A All localities. B Gara Sbaa. X-axis indicates size in 20 mm size bins; Y-axis indicates numbers of specimens. Specimen size bins: 1 0-20 mm 2 21-40 mm 3 41-60 etc.
Figure 48.
Figure 48.
Size of vertebrate fossil elements found in the Kem Kem Group (FSAC-KK collection). A Boumerade. B Aferdou N’Chaft. X-axis indicates size in 20 mm size bins; Y-axis indicates numbers of specimens. Specimen size bins: 1 0-20 mm 2 21-40 mm 3 41-60 etc.
Figure 49.
Figure 49.
Size of vertebrate fossil elements found in the Kem Kem Group. A Iferda N’Ahouar B 'Valley near Boumerade' locality. X-axis indicates size in 20 mm size bins; Y-axis indicates numbers of specimens. Specimen size bins: 1 0-20 mm 2 21-40 mm 3 41-60 etc.
Figure 50.
Figure 50.
Possible biofilm (bacteria, fungi) from Oum Tkout, Douira Formation. Scale bar equals 5 mm.
Figure 51.
Figure 51.
Simple and compound leaves are among the plant remains from Oum Tkout, Douira Formation. Scale bars equal 1 cm in A and B, 5 mm in C and 5 cm in D.
Figure 52.
Figure 52.
Decapod prawn Cretapenaeus berberus (Garassino et al. 2006) from Oum Tkout, Douira Formation. Scale bar equals 5 mm.
Figure 53.
Figure 53.
Dragonfly larva (Odonata indet.) from Oum Tkout, Douira Formation. Scale bar equals 5 mm.
Figure 54.
Figure 54.
Selected isolated remains of the sclerorhynchid Onchopristis numidus. Rostral fragment with the bases of two rostral teeth in place (NMC 41876) in (A) ?dorsal, (B) ?ventral and (C) lateral view. Section of a large rostrum (FSAC-KK 937) in (D) dorsal, (E) ventral, (F) anterior and (G) posterior views H Small section of rostrum (NMC 50397) I Isolated rostral tooth J Placoid scale (MNHN-MRS 72) D and H reconstructed to show the approximate shape of the rostral blade. Scale bars equal 5 cm in A-H, 2 cm in I and J.
Figure 55.
Figure 55.
Elasmobranch fossils from Morocco and Egypt. Hybodontoidea fin spine with tubercles (FSAC-KK 943), in (A) lateral and (B) posterior view. Hybodontoidea fin spine (FSAC-KK 944), ornamented with striations in (C) lateral and (D) posterior view. Lamniform tooth (Poi-SGM 52) referred to Haimirichia amonensis in (E) labial and (F) lingual view. Lamniform tooth (Poi-SGM 53) referred to Cenocarcharias cf. tenuiplicatus in (G) labial and (H) lingual views. I Rostral tooth of Peyeria libyca (Poi SGM 61). Scale bars equal 3 cm in A-D, 3 mm in E and F, 2 mm in G and H, 1 cm in I.
Figure 56.
Figure 56.
Associated elasmobranch vertebrae likely referable to Onchopristis numidus from the Kem Kem Group. A Reconstructed vertebral series. B-D Select vertebrae with sedimentary matrix. Scale bars equal 10 cm in A, 5 cm in B-D.
Figure 57.
Figure 57.
Serenoichthys kemkemensis Dutheil, 1999 from the Douira Formation. Scale bar equals 1 cm.
Figure 58.
Figure 58.
Large polypterid premaxilla (FSAC-KK 209) from the Kem Kem Group. In (A) anterior, (B) posterior, (C) lateral, (D) dorsal and (E) ventral view. Scale bar equals 3 cm. Abbreviations: f foramen t tooth.
Figure 59.
Figure 59.
Large polypterid (MNBH IGU23) from the Cenomanian of Niger. Associated right and left premaxillae in (A) anterior, (B) ventral and (C) lingual view. Scale bar equals 3 cm.
Figure 60.
Figure 60.
Scales of a large holostean from the Kem Kem Group. AFSAC-KK 530 (Gara Sbaa) B NMC 50434-A CFSAC-KK 531 (Gara Sbaa) D NMC 50434-C E NMC 50434-B F Lateral view of NMC 50817 G Medial view of NMC 50817 H Lateral view of NMC 41932 I Medial view of NMC 41932 (one of the largest known teleost scales from the Kem Kem Group). Scale bar equals 3 cm.
Figure 61.
Figure 61.
Actinopterygian remains from the Kem Kem Group. A-C Possible amiid dentary fragment (MDM 02) in right lateral, medial and dorsal (occlusal) views D Possible amiid dentary fragment (MSNM V 6417) E Dentary of an unidentified teleost (NMC 41883) F Possible notopterid dental plate (NMC 50863) G Probable teleost bone (NMC 41900) H Possible teleost parasphenoid (NMC 50864) I, J Possible Obaichthys africanus (Cavion et al. 2015) scales (NMC 50437) K, L Possible seminotiform scale (NMC 50867A) M-P Possible lepisosteid or obaichthyid vertebra in anterior, posterior, dorsal and ventral view. Scale bars equal 1 cm in A-D, 2 cm in E-L, 2 cm in M-P.
Figure 62.
Figure 62.
Isolated teleost dentaries from the Kem Kem Group. A-C Partial dentary of an ichthyodectiform with affinities to Xiphactinus and Cladocyclus (NMC 41882) in right lateral, medial and dorsal views D Right dentary of Xiphactinus audax (FHSM VP-2973) from the Late Cretaceous of the USA in medial view (courtesy of Mike Everhart) E-I Isolated dentary (FSAC-KK 906) of possible amiid in right lateral, medial, dorsal (occlusal) ventral, and medial (close-up) views J-K Dentary (NMC 41884) in ?right lateral and dorsal (occlusal) view. L-M Dentary of an unidentified predatory teleost with rostral tooth of Onchopristis numidus (NMC 50836) in lateral and medial view. Scale bars equal 5 cm in A-C, E-H, 10 cm in D, 3 cm in I-K, 5 cm in L and M. Abbreviation: rt rostral tooth.
Figure 63.
Figure 63.
Diplospondichthys moreaui Filleul and Dutheil, 2004 from the Kem Kem Group. Scale bar equals 2 cm.
Figure 64.
Figure 64.
Spinocaudichthys oumtkoutensis Filleul and Dutheil, 2002 from the Kem Kem Group. Scale bar equals 5 mm.
Figure 65.
Figure 65.
Isolated elements referable to Axelrodichthys lavocati or a closely related form from the Kem Kem Group. A Isolated pterotic (NMC 41877) B Right angular (NMC 50816). Parasphenoid (NMC 41813) in (C) dorsal, (D) anterior and (E) posterior view. Visceral arch element (NMC 50828) in (F) lateral and (G) medial view. Visceral arch element (NMC 50827) in (H) lateral and (I) medial view. Isolated postparietal of (FSAC-KK 157), collected at locality 5 (Figure 9) in (J) dorsal and (K) ventral views. L Isolated operculum (MNHN-MRS 926). M Left angular (MNHN-MRS 78), part of type material of A. lavocati (Tabaste 1963). Scale bars equal 5 cm in A–I, 5 cm in J and K, 10 cm in L and M. Abbreviations: ant ap anterior apophysis desc pr descending process of postparietal.
Figure 66.
Figure 66.
Large specimens from the Kem Kem Group likely referable to Axelrodichthys. Possible fragment of palatoquadrate region preserving a large part of the pterygoid (MNHN-MRS 926) in (A) lateral and (B) medial view. Large quadrate with a small section of the pterygoid (MPDM 14) in (C) lateral and (D) medial view. E Partial palate of Axelrodichthys (MNHN-MRS 1761) in lateral view. Quadrate (NMC 41994) in (F) lateral, (G) medial and (H) ventral view. Scale bars equal 10 cm in A, B and E, and 3 cm in C, D and F-H. Abbreviations: pt pterygoid q quadrate.
Figure 67.
Figure 67.
Size variation in lungfish tooth plates from the Kem Kem Group. AUCRC PV62, likely Neoceratodus africanusBFSAC-KK 2735. Note that some size differences can be even greater in exceptional specimens. Scale bar equals 3 cm.
Figure 68.
Figure 68.
Amphibia and Squamata from the Douira Formation. Cf. Kababisha sp. braincase (UCRC PV50) from Oum Tkout in (A) lateral and (B) posterior view. Oumtkoutia anae braincase (UCRC PV63) from Oum Tkout in (C) lateral and (D) ventral view. Simoliophis cf. S. lybicus trunk vertebra UCRC PV127) from Douira in (E) anterior and (F) lateral view. Figures modified from Rage and Dutheil (2008). Scale bars equal 1 mm in A and B, 2 mm in C and D, 3 mm in E and F.
Figure 69.
Figure 69.
Testudinid shell fragments from the Kem Kem Group. Partial plastron with sutures (UCRC PV165) in (A) dorsal and (B) ventral view. Large shell fragment (UCRC PV166) with sutures highlighted by red and dark lines (C). Rounded and smooth shell fragment (MNHN-MRS 290) (D). MNHN-MRS 382 (E). Dark-colored carapace elements from Kouah Trick locality (MNHN-MRS 1813-1815) in (F, H, J) dorsal and (G, I, K) ventral view. Scale bar equals 5 cm.
Figure 70.
Figure 70.
Possible araripemydid and podocnemidinuran shell fragments from the Kem Kem Group. Possible araripemydid plastral element (MNHN-MRS 172) in (A) dorsal and (B) ventral view. MNHN-MRS 1175 in (C) dorsal and (D) ventral view. NMC 41928 in (E) dorsal and (F) ventral view. Partial podocnemidinuran (?Galianemys) plastron collected at Iferda N’Ahouar (UCRC PV167) in (G) dorsal and (H) ventral view. Cross-sectional view of a typical shell fragment showing dense and cancellous bone (I). Scale bars equal 3 cm in A-F, 3 cm in G and H, 1 cm in I.
Figure 71.
Figure 71.
Isolated turtle postcrania from the Kem Kem Group. Pelomedusoides indet., cervical vertebra (BSPG 2006 I 61) in (A) dorsal, (B) left lateral, (C) anterior and (D) posterior view. ?Testudines indet., proximal part of femur (NMC 41973) in (E) anterior and (F) posterior view. Testudines indet., metapodial (NMC 41975) in (G) anterior, (H) proximal and (I) distal view. Testudines indet., selected metapodials (NMC 50846 A-E) in (J) dorsal view. Scale bars equal 3 cm in A-D, 3 cm in E and F, 3 cm in G-J.
Figure 72.
Figure 72.
Partial shell and girdle bones of cf. Galianemys sp. (UCRC PV18) in (A) dorsal and (B) anterodorsolateral view. Scale bar for A: 10 cm.
Figure 73.
Figure 73.
Turtle skull elements from the Kem Kem Group. One of two undescribed skulls likely referable to Galianemys whitei (MNHN-MRS 2098) in (A) dorsal, (B) ventral, (C) anterior and (D) right lateral view. Partial skull of Galianemys sp. (FSAC-KK 938) in (E) dorsal, (F) ventral and (G) anterior view. Skull of Dirqadim schaefferi (MDEt 41) in (H) dorsal, (I) ventral and (J) right lateral view. Scale bars equal 3 cm in A-D, E-G and H-J. Abbreviations: bo basioccipital, f frontal, m maxilla, op opisthotic, p parietal, pf prefrontal, pl palatine, pt pterygoid, q quadrate, so supraoccipital.
Figure 74.
Figure 74.
Specimens referred to Araripesuchus rattoides from the Kem Kem Group. NMC 41893 in (A) right lateral, (B) medial, (C) dorsal (occlusal) and (D) anterior view. BSPG 2008 I 41 in (E) left lateral and (F) dorsomedial view. Scale bars equal 3 cm in A-C, E and F, and 1 cm in D.
Figure 75.
Figure 75.
Sebecid skull, referred to Hamadasuchus rebouli (ROM 52620) in (A) left lateral, (B) dorsal and (C) ventral views (from H.-D. Sues; Larsson and Sues 2002). Scale bar equals 10 cm. Abbreviations: f frontal m maxilla n nasal pl palatine pm premaxilla pt pterygoid q quadrate qj quadratojugal.
Figure 76.
Figure 76.
Other sebecid cranial specimens. Posterolateral skull roof (FSAC-KK 930) in (A) dorsal and (B) right lateral view. Partial braincase of Hamadasuchus rebouli (MNHN-MRS 3101) in (C) dorsal view (initially referred to Libycosuchus, Buffetaut 1994). Scale bars equal 10 cm in A and B, 5 cm in C.
Figure 77.
Figure 77.
Original material of Libycosuchus brevirostris Stromer, 1914 (BSP 1912 VIII 574-578). Cranium in (A) dorsal, (B) ventral (C) left lateral view. Lower jaw in (D) dorsal and (E) ventral view. Skull in (F) dorsolateral view. ?Sacral vertebra in (G) lateral view. Caudal vertebra in (H) anterior and (I) lateral view. Scale bars equal 10 cm in A-E, 3 cm in G-I.
Figure 78.
Figure 78.
Comparison of the braincases – in dorsal view – of the small crocodyliforms (A) Libycosuchus brevirostris, (B) sebecid indet. (FSAC-KK 08) and (C) a specimen initially referred to Hamadasuchus rebouli (MNHN-MRS 3101). Specimens adjusted to similar size (see Figs 76, 77 and 81 for scale in A-C).
Figure 79.
Figure 79.
Skull of Hamadasuchus rebouli (BSPG 2005 I 83, Rauhut & López-Arbarello, 2006) in (A) lateral and (B) dorsal view (see). Scale bar equals 5 cm.
Figure 80.
Figure 80.
Sebecid upper jaw bones from the Kem Kem Group. Partial left maxilla (NMC 41866) in (A) lateral and (B) medial view. Partial left premaxilla (NMC 41892) in (C) anterolateral and (D) medial view. Maxillary fragment (FSAC-KK 932) in (E) lateral, (F) medial and (G) ?anterior view. Base of the maxillary crown (FSAC-KK 932) in (H) lateral view. Scale bar equals 3 cm in A-G, 1 cm in H.
Figure 81.
Figure 81.
Sebecid braincases from Aferdou N’Chaft. Braincase (FSAC-KK 1237) referable to Hamadasuchus rebouli in (A) dorsal and (B) ventral view. Partial braincase (FSAC-KK 08) in (C) dorsal, (D) ventral, (E, I) anterior, (F) posterior, (G) left lateral and (H) right lateral view. Scale bar equals 5 cm in A-H. Abbreviations: bo basioccipital bsr basisphenoid recess cqp cranioquadrate passage f frontal fo ICA foramen for internal carotid artery fo MM maxillomandibular foramen fo mAMEP Musculus adductor mandibulae externus profundus fo V1 ophthalmic foramen oc occipital condyle p parietal q quadrate.
Figure 82.
Figure 82.
Mandibular symphyses of Hamadasuchus rebouli. Mandibular symphysis (MNHN-MRS 3110) in (A) dorsal, (B) ventral and (C) right lateral view. Mandibular symphysis (NMC 41784) in (D) dorsal, (E) ventral, (F) left lateral, (G) anterior and (H) posterior view. Scale bar equals 3 cm. Abbreviations: ad2, 4 alveolus for dentary tooth 2, 4 d4 dentary tooth 4 fo foramen sp splenial.
Figure 83.
Figure 83.
Specimens of Laganosuchus from the Echkar Formation in Niger and the Kem Kem Group in Morocco. Cast of lower jaws (UCRC PVC9) of the holotypic specimen of L. thaumastos (MNN IGU13) in (A) dorsal and (B) left lateral view. Holotypic specimen of L. maghrebensis (UCRC PV2) from the Kem Kem Group in (C) left lateral, (D) dorsal and (E) ventral view. BSPG 2008 I 62 in (F) left lateral, (G) dorsal (occlusal) and (H) ventral view. Partial dentary of L. maghrebensis (NMC 50838) in (I) lateral, (J) dorsal (occlusal) and (K) ventral view. Scale bars equal 20 cm in A and B, 6 cm in C–K.
Figure 84.
Figure 84.
Dentary fragment with tooth referred to Laganosuchus maghrebensis (NMC 41786) in (A) lateral, (B) medial and (C) dorsal view. Scale bar equals 2 cm. Abbreviation: d dentary.
Figure 85.
Figure 85.
Holotypic specimen of Aegyptosuchus peyeri Stromer, 1933 from the Bahariya Formation of Egypt in dorsal (A) and ventral (B) view. Scale bar equals 10 cm.
Figure 86.
Figure 86.
Cranial remains of the crocodyliform Elosuchus. A, CE. cherifiensis cranium (MNHN SAM 129) in dorsal and ventral views BE. cherifiensis (MNHN SAM 129) with posterior skull roof (NMC 41912) and anterior snout section (FSAC-KK 10) superimposed, DE. cherifiensis premaxilla (MNHN SAM 129), drawing in ventral view EE. cherifiensis premaxilla FSAC-KK 10 FE. cherifiensis occipital piece (MNHN-MRS 340-25) GE. cherifiensis lower jaw reconstructed from several specimens (MNHN-E 43, SAM 138, SAM 137-157; redrawn from Lapparent de Broin 2002) HE. cherifiensis anterior dentary symphysis (MNHN E 43) in dorsal view I Partial dentary symphysis (MNHN-INA 25). Abbreviations: bo basioccipital, d dentary, ex, exoccipital j jugal m maxilla pm premaxilla pt pterygoid sym symphysis. Scale bars equal 10 cm in A-C and E-G, 20 cm in H.
Figure 87.
Figure 87.
Rostral fragments referred to cf. Elosuchus. Rostral fragment (FSAC-KK 10) in (A) dorsal, (B) ventral, (C) right lateral and (D) anterior view. Rostral fragment (NMC 41866) in (E) dorsal and (F) ventral view. Maxillary piece (MNHN-MRS 3111) in (G) dorsal and (H) ventral view. Scale bar equals 10 cm.
Figure 88.
Figure 88.
Braincase and jugal of the crocodyliform Elosuchus cherifiensis. Braincase (MNHN-MRS 3115) in (A) dorsal and (B) ventral view. Braincase (NMC 41912) in (C) dorsal, (D) ventral and (E) left lateral view. Jugal (FSAC-KK 09) in (F) left lateral, (G) medial and (H) ventral view. Scale bar equals 20 cm.
Figure 89.
Figure 89.
Fragmentary bones tentatively referred to the crocodyliform Elosuchus. Left premaxilla (FSAC-KK 923) in (A) lateral, (B) medial, (C) anterior, (D) dorsal and (E) ventral view. ?Lacrimal (BSPG 2008 I 60) in (F) lateral and (G) medial view. Basioccipital and basisphenoid (FSAC-KK 926) in (H) ventral and (I) posterior view. Left dentary fragment (UCRC PV168) in (J) dorsal view. K Left dentary fragment with five teeth (NMC 41785) in lateral view. Scale bar equals 5 cm. Abbreviations: bo basioccipital pm4 premaxillary tooth 4.
Figure 90.
Figure 90.
Dentaries of the crocodyliform Elosuchus cherifiensis. A Left anterior dentary (UCRC PV169) in dorsal view B Dentary symphysis (E 44 (MNHN; Lapparent de Broin 2002) in dorsal view C, D Left anterior dentary (NMC 41867) in dorsal and left lateral views E-H Dentary symphysis (NMC 41791) in dorsal, ventral, right lateral and anterior view. I, J Dentary and splenial symphysis (MNHN-MRS 3112) in dorsal and ventral view. Scale bars equal 10 cm. Abbreviations: pm1 premaxillary tooth 1 sp splenial.
Figure 91.
Figure 91.
Mandibular symphysis of the crocodyliform Elosuchus cherifiensis. Right anterior dentary (FSAC-KK 753) in (A) dorsal, (B) ventral, (C) right lateral, (D) medial and (E) anterior view. Reconstructed symphysis of Elosuchus cherifiensis using reflected image of FSAC-KK 753 in (F) dorsal and (G) ventral view. Scale bar equals 5 cm in A-D, F and G, and 2 cm in E. Abbreviations: d6, 12 dentary tooth 6, 12 sp splenial.
Figure 92.
Figure 92.
Isolated pholidosaurid crocodyliform teeth. A NMC 41791 (see Fig. 88E) B-DBSPG 1993 IX 334 EMNHN-MRS 765 FMNHN-MRS 766 GMNHN-MRS 767. Scale bar equals 3 cm.
Figure 93.
Figure 93.
Sebecid left jugal (FSAC-KK 07) in (A) lateral, (B, D) medial and (C) ventral view. Scale bar equals 5 cm in A–C. Abbreviations: nAD dorsal alveolar branch of the maxillary nerve (V2) nJU jugal branch of the maxillary nerve (V2).
Figure 94.
Figure 94.
Ornithocheirid pterosaur teeth from the Kem Kem Group. AFSAC-KK 885 BFSAC-KK 44 CFSAC-KK 886 D, EFSAC-KK 197 in labial and lingual views FFSAC-KK 887 GFSAC-KK 941 H Teeth of ‘morphotype 1’ of Wellnhofer and Buffetaut (1999) (BSPG 1993 IX 590-596) I Teeth of ‘morphotype 2’ of Wellnhofer and Buffetaut (1999) (BSPG 1993 IX 597-607) J Teeth of ‘morphotype 3’ of Wellnhofer and Buffetaut (1999) (BSPG 1993 IX 608-617) K Tooth of ‘morphotype 4’ of Wellnhofer and Buffetaut (1999) (BSPG 1993 IX 618). Scale bar equals 3 cm.
Figure 95.
Figure 95.
Ornithocheirid pterosaur jaw fragments from the Kem Kem Group. Premaxillae of Siroccopteryx moroccensis in (A) left lateral and (B) ventral views (from Mader and Kellner 1999). Dentary ramus of an indeterminate ornithocheirid (FSAC-KK 33) in (C) ?left lateral and (D) ?dorsal view. Scale bars equal 3 cm. Abbreviations: ad alveolus for dentary tooth iap interalveolar pit pm2 premaxillary tooth 2.
Figure 96.
Figure 96.
Rostral fragment (BSP 1993 IX 338) referred by McPhee et al. (2020) to a new ?chaoyangopterid azhdarchoid from the Kem Kem Group in (A) left lateral, (B) right lateral, (C) dorsal, (D) ventral and (E) posterior view. Fragmentary azhdarchoid rostrum (FSAC-KK 27) in (F) left lateral, (G) right lateral, (H) dorsal, (I) ventral and (J) posterior view. Scale bars equal 5 cm in A-E, 3 cm in F-J.
Figure 97.
Figure 97.
Fragment of the mandibular symphysis of a tapejarid pterosaur (BSP 1997 I 67) from the Kem Kem Group in (A) ?left lateral, (B) ?right lateral and (C) dorsal (occlusal) view. Scale bar equals 5 cm.
Figure 98.
Figure 98.
Rostrum of Alanqa saharica (FSAC-KK 26) from the Kem Kem Group in (A) ventral and (B) lateral views with magnified view of foramina and ventral margin. Scale bar equals 5 cm.
Figure 99.
Figure 99.
Fragment, coated in ammonium chloride, of the mandibular symphysis of Xericeps curvirostris, Martill et al. 2018, FSAC-KK 10700 in (A) left lateral, (B), occlusal, (C), right lateral and (D), ventral views. Reproduced from Martill et al. 2018. Abbreviations: os occlusal surface aos accessory occlusal surface l fo lateral foramina vs ventral sulcus. Scale bar equals 50 mm.
Figure 100.
Figure 100.
Fragments of jaws of a ?chaoyangopterid pterosaur from the Kem Kem Group. FSAC-KK 29 in (A) left lateral, (B) ?dorsal, (C) anterior and (D) posterior views E Detailed view of anterior-most paired foramina. FSAC-KK 32 in (F) left lateral, (G) ?dorsal and (H) posterior view. UCRC PV161 in (I) left lateral, (J) ?dorsal and (K) posterior view. Scale bar equals 5 cm in A-D and 4 cm in F-K. E scale bar equals 5 mm.
Figure 101.
Figure 101.
Azhdarchid pterosaur rostral fragment from the Kem Kem Group. FSAC-KK 28 in (A) lateral, (B) ?dorsal, (C) ?ventral and (D) posterior view. Scale bar equals 3 cm.
Figure 102.
Figure 102.
Near complete ?third cervical of an azhdarchid pterosaur from the Kem Kem Group. FSAC-KK 3088 in (A) ventral, (B) dorsal, (C) right lateral, (D) left lateral, (E) anterior and (F) posterior view. Scale bar equals 5 cm.
Figure 103.
Figure 103.
Possible thyreophoran tooth. UCRC PV162 unnumbered in (A) labial and (B) lingual view. Scale bar equals ~5 mm. Abbreviations: cr crown dtl denticles.
Figure 104.
Figure 104.
Holotype material of Rebbachisaurus garasbae. A Dorsal vertebra (MNHN-MRS 1957) in anterior view and (B) lateral view. C Scapula (MNHN-MRS 1957) in lateral view. Scale bar equals 30 cm. Abbreviations: ac fo acromial fossa ac no acromial notch nc neural canal med spol medial spinopostzygapophyseal lamina.
Figure 105.
Figure 105.
Left humerus of Rebbachisaurus garasbae. MNHN-MRS 2002 in (A) anterior, (B) posterior, (C) lateral and (D) cross-sectional view. Scale bar equals 20 cm. Abbreviations: dpc deltopectoral crest hdr ridge emanating from humeral head schc insertion for M. scapulo-humeralis cranialis.
Figure 106.
Figure 106.
Axial remains of Rebbachisaurus garasbae. NMC 50844 in (A) anterior, (B) posterior, (C) right lateral and (D) dorsal views E Internal appearance of centrum F Posterior articular section of vertebra (MNHN-MRS 1184) G Partial centrum (MNHN-MRS 1857) in right lateral view. Scale bar equals 20 cm. Abbreviations: c centrum cpol centropostzygapophyseal lamina lat. cprl lateral centroprezygapophyseal lamina nc neural canal pc pleurocoel poz postzygapophysis.
Figure 107.
Figure 107.
Sauropod tooth collected at Aferdou N’Chaft, tentatively referred to Rebbachisauridae. FSAC-KK 910 in (A) labial and (B) lingual view. Scale bar equals 3 cm. Abbreviation: wf wear facet.
Figure 108.
Figure 108.
Isolated sauropod teeth from the Kem Kem Group. A NMC 50822 in lingual view B NMC 50825 in labial view C NMC 41808 in labial view. NMC 41812 in (D) labial and (E) lingual views F NMC 41809 in labial view G NMC 50442B in labial view H NMC 41801 in labial view I NMC 41799 in lingual view J NMC 41811 in lateral view. NMC 41810 in (K) lingual and (L) cross-sectional view. NMC 50445 in (M) cross-sectional, (N) labial and (O) lingual views P NMC 50442A in labial view Q-SBSPG 1993 IX 331 in (Q) lateral and (R, S) lingual views respectively T NMC 50442D in lingual view U NMC 41810 in labial view V NMC 50824 in labial view W NMC 50442 in labial view. Scale bar equals 3 cm.
Figure 109.
Figure 109.
Proximal section of a right ulna of a giant sauropod. UCRC PV9 in (A) anterior, (B) posterior and (C) distal view. Scale bar equals 20 cm. Abbreviations: alp anterolateral process amp anteromedial process opr olecranon process.
Figure 110.
Figure 110.
Indeterminate theropod teeth from the Kem Kem Group. MNHN-MRS 783 in (A) lateral and (B) cross-sectional views CFSAC-KK 914 in lateral view DMNHN-MRS 1457 in lateral view EFSAC-KK 913 in lateral view. FSAC-KK 909 in (F) labial and (G) lingual views H Detailed view of FSAC-KK 909. IFSAC-KK unnumbered in lateral view J NMC 50446-C KFSAC-KK 915 in lateral view LFSAC-KK 912 in lateral view MFSAC-KK 916 in labial view N Detailed view of denticles of FSAC-KK 916. Scale bar equals 2 cm in A-C, E-G and I-M, 3 cm in D and 5 mm in H and N.
Figure 111.
Figure 111.
Isolated unguals from the Kem Kem Group. NMC 41820 in (A) posterior and (B) lateral views C NMC 50987 (cast) in lateral view. NMC 50842b in (D) posterior and (E) lateral view. FSAC-KK 918 in (F) lateral, (G) dorsal and (H) ventral views I NMC 50839a in lateral view J NMC 50839b in lateral view K 50839c (reversed) in lateral view L NMC 50839e in lateral view M NMC 50839d (reversed) in lateral view NMPDM 43 in lateral view OBSPG 1993 IX 311 in lateral view PBSPG 1993 IX 330 in lateral view. NMC 50826 in (Q) posterior and (R) lateral views S NMC 50386 in lateral view. NMC 50842A in (T) posterior and (U) lateral views VBSPG 1993 IX 330 in lateral view WFSAC-KK 202 in lateral view. FSAC-KK 917 in (X) posterior and (Y) lateral view. Scale bar equals 5 cm. Abbreviations: ftu flexor tubercle lg lateral groove.
Figure 112.
Figure 112.
Abelisaurid jaw elements and teeth from the Kem Kem Group. Right maxilla (UCPC PV10) in (A) lateral, (B) medial, (C) anterior and (D) dorsal view. ?Abelisaurid tooth (MSNM V6053) in (E) labial view. ?Abelisaurid tooth (NMC 50446) in (F) labial view. Right dentary section (NMC 41861) in (G) lateral and (H) medial view. Scale bars equal 5 cm in A-D, G and H, 2 cm in E and F.
Figure 113.
Figure 113.
Abelisaurid maxillae from Madagascar and Niger. A Left maxilla of Majungasaurus crenatissimus (FMNH PR 2100, cast) in lateral view B Left maxilla of Kryptops palaios (MNN GAD1) C Reconstructed skull of Rugops primus (modified from Sereno et al. 2004) with maxilla in gray. D, E Left maxilla of Rugops primus (MNN IGU1) in lateral and medial view. Scale bar equals 10 cm in A, B, D and E.
Figure 114.
Figure 114.
Comparison of abelisaurid maxillae from Morocco and Niger. A Left maxilla (UCPC 10, mirrored) in ventral view B Left maxilla of Rugops primus (MNN IGU1) in ventral view. Scale bar equals 5 cm.
Figure 115.
Figure 115.
Excavation and skeletal reconstruction of Deltadromeus agilis. A Excavation of the fibula B Mounted skeletal reconstruction using casts of the holotype UCRC PV11. Scale bar equals 5 cm in A. Length of skeleton in B is about 8 m.
Figure 116.
Figure 116.
Well-preserved elements of the right pes of D. agilis. A Digit IV in dorsal view. B Digit III in dorsal view. C Digit II in dorsal view. Scale bar equals 10 cm. Abbreviations: mt II, III metatarsals II, III ph III-1, 2 pedal phalanges 1 and 2 of digit III ph IV-3, 4, 5 pedal phalanges 3-5 of digit IV.
Figure 117.
Figure 117.
Left metatarsals of Deltadromeus agilis (UCRC PV11). A Lateral view B Detailed view of the first phalanx of digit IV. Scale bar equals 10 cm in A. Abbreviations: mt III, IV metatarsal III, IV ph IV-1 phalanx 1 of digit IV.
Figure 118.
Figure 118.
Deltadromeus agilis from Morocco and Egypt. A Revised reconstruction based on UCRC PV11 B Holotype compared to a large femur (now lost) referred to the genus and species from the Bahariya Formation, Egypt. Known elements in white. Scale bars: 1 m.
Figure 119.
Figure 119.
Tooth of Spinosaurus aegyptiacus (FSAC-KK 08) from the Kem Kem Group in (A) side, (B) opposing side and (C) basal view. Scale bar equals 5 cm.
Figure 120.
Figure 120.
Dentary fragments of Spinosaurus aegyptiacus from the Kem Kem Group. Dentary section (MPDM 30) in (A) lateral and (B) ?anterior view. Dentary fragment (MNHN-MRS 1513) in (C) lateral and (D) dorsal view. Left dentary fragment with tooth (MSNM V6865) in (E) lateral view. Dentary fragment (MPDM 31, associated with Onchopristis rostral tooth) in (F) dorsal view. Scale bar equals 10 cm. Abbreviations: f foramen rt rostral tooth.
Figure 121.
Figure 121.
Dentary fragments of Spinosaurus aegyptiacus from the Kem Kem Group. Anterior section of the left dentary (NMC 50833) in (A) left lateral, (B) medial, (C) dorsal, (D) ventral, (E) anterior and (F) posterior view. Symphyseal end of the right dentary (NMC 50832) in (G) lateral, (H) medial, (I) dorsal, (J) ventral, (K) anterior and (L) posterior view. Scale bar equals 10 cm. Abbreviations: ad1 3 alveolus for dentary tooth 1, 3 dl dental lamina Mc Meckel’s canal t tooth.
Figure 122.
Figure 122.
Dentary fragments of Spinosaurus aegyptiacus from the Kem Kem Group. Anterior section of the left dentary (IMPG 969-1) in (A) lateral and (B) dorsal view. Posterior section of the left dentary (IMPG 969-2) in (C) lateral and (D) medial view. Part and counterpart aligned in (E) lateral and (F) dorsal view. The two specimens clearly belong to the same jaw section and individual (contraBuffetaut 1989, 1992). Scale bar equals 10 cm in A-D. Abbreviations: ad alveolus in the dentary t tooth.
Figure 123.
Figure 123.
Comparison of Spinosaurus aegyptiacus rostra from Morocco and Algeria. Moroccan rostra include MSNM V4047 in (A) left lateral view and BMNH 16420 in (B) left lateral view. Algerian rostrum MNHN SAM 124 is shown in (C) dorsal, (D) ventral and (E) anterior view. Scale bars equal 20 cm in A-D, 10 cm in E.
Figure 124.
Figure 124.
Rostral portions of Spinosaurus from Morocco and Algeria. Rostrum (MSNM V4047) in (A) left lateral view with enlarged sections. Rostrum (MNHN SAM 124) from Algeria in (B) ventral view with enlarged sections. Scale bar equals 20 cm for full lateral and ventral view. Abbreviations: antfe antorbital fenestra bfr bone fractures en external naris f foramen m2, 4 maxillary tooth 2, 4 m maxilla n nasal pm premaxilla rpm rostro-medial process of maxillae sf subnarial foramen.
Figure 125.
Figure 125.
Dentary portion of Spinosaurus aegyptiacus from the Kem Kem Group. NHMUK VP 16421 in (A) left lateral, (B) medial, (C) dorsal (occlusal), (D) anterior and (E) posterior view. F Detailed view of dentary alveolus. Size comparison of (G) the anterior end of the dentary of Spinosaurus aegyptiacus (NHMUK VP 16421) and (H) the partial dentary of Baryonyx walkeri (NHMUK VP R9951). Scale bars equal 20 cm in A-E, 10 cm in G and H.
Figure 126.
Figure 126.
Cervical vertebrae of Spinosaurus cf. aegyptiacus from the Kem Kem Group. Mid cervical vertebra (NMC 50791) in (A) right lateral, (B) anterior, (C) posterior, (D) dorsal and (E) ventral view. Posterior cervical vertebra (NMC 50790) in (F) right lateral, (G) anterior and (H) ventral view. Mid cervical vertebra (NMC 41768) in (I) left lateral view. Posterior cervical vertebra (MPDM 33) in (J) left lateral and (K) dorsal view. Scale bar equals 20 cm. Abbreviations: di diapophysis epi epipophysis ila interspinous ligament attachment ns neural spine pa parapophysis pc pleurocoel prz prezygapophysis.
Figure 127.
Figure 127.
Dorsal vertebrae of Spinosaurus aegyptiacus. Anterior dorsal neural arch with partial spine (FSAC-KK 04) from the Kem Kem Group in (A) left lateral, (B) right lateral, (C) anterior and (D) posterior view. Ventral portion of a mid dorsal vertebra (Stromer 1915) from the Bahariya Formation in Egypt in (E) right lateral view. Scale bar equals 10 cm in A-D. Abbreviations: c centrum id fos infradiapophysial fossa ipoz fos infrapostzygapophysial fossa iprz fos infraprezygapophysial fossa ns neural spine.
Figure 128.
Figure 128.
Problematic cervical vertebrae from Niger and Morocco referable to Spinosaurus. NMC 41629 in (A) right lateral, (B) ventral, (C) anterior and (D) posterior view. NMC 41774 in (E) anterior and (F) posterior view. MNN IGU 11 in (G) right lateral, (H) ventral and (I) posterior view. UCRC PV13 in (J) anterior and (K) posterior view. NMC 41857 in (L) dorsal, (M) anterior and (N) posterior view. Scale bar equals 10 cm. Abbreviations: di diapophysis k keel pa parapophysis prz prezygapophysis.
Figure 129.
Figure 129.
Skull and skeleton of Spinosaurus aegyptiacus. A Skull reconstruction in left lateral view (after Dal Sasso et al. 2005) B Skull reconstruction with adult rostrum (MSNM V4047) and dentary (NHMUK VP R 16421) superimposed C Skeletal reconstruction (swimming pose, after Ibrahim et al. 2014b). Scale bars equal 40 cm in A, B and 1 m in C. Abbreviations: a angular d dentary, f frontal, j jugal m maxilla n nasal pm premaxilla po postorbital qj quadratojugal sa, surangular.
Figure 130.
Figure 130.
Spinosaurid cervicodorsal vertebrae. A Partial mid cervical of Spinosaurus aegyptiacus (FSAC-KK 11888, cast) lacking the anterior end of the centrum, upper portion of the neural spine and postzygapophyses, B Articulated cervicodorsal series C2-D4) of Suchomimus tenerensis (MNBH GAD70) in left lateral view with most of the cervical ribs removed. Scale bar equals 5 cm in A (bottom), 10 cm in A (top) and B. Abbreviations: ns neural spine pl pleurocoel plf platform prz prezygapophysis tp transverse process.
Figure 131.
Figure 131.
Teeth from Morocco and Niger referred to Carcharodontosaurus. Teeth of UCRC PV12 (C. saharicus) in (A) labial and (B) lingual view. C Detail of wrinkled enamel D Cross-sectional view. EMPDM 16 in lingual view. F Tooth (UCRC PV163) in lingual view. G Tooth (UCRC PV164) in lingual view. HFSAC-KK 907 in labial view. Scale bars equal 5 cm in A and B, and E-H, 1 cm in C and 3 cm in D. Abbreviation: wr wrinkled enamel.
Figure 132.
Figure 132.
Endocasts of Carcharodontosaurus saharicus. Endocast (from UCRC PV12) in (A) left lateral, (B) right lateral, (C) dorsal and (D) ventral view. Endocast (MB. R. 2056) in (E) dorsal, (F) right lateral views and (G) bottom of cast. Scale bars equal 10 cm in A-D, 5 cm in E and F. Abbreviations: cer cerebrum ls longitudinal sinus vcd vena capitus dorsalis I, V cranial nerves I and V.
Figure 133.
Figure 133.
Right maxilla of Carcharodontosaurus saharicus. UCRC PV12 in (A) lateral and (B) medial view. (See also Brusatte and Sereno 2007). Scale bar equals 20 cm. Abbreviations: amp anteromedial process antfor antorbital fossa ridge gdl groove for dental lamina iad interalveolar depression ip interdental plate m maxilla m3, 6 maxillary teeth 3, 6 mf maxillary fenestra.
Figure 134.
Figure 134.
Nasal and lacrimal of Carcharodontosaurus saharicus (UCRC PV12). Right nasal in (A) lateral and (B) medial views C Left nasal in medial view. Left lacrimal in (D) lateral and (E) medial view. Scale bar equals 20 cm. Abbreviations: lfo lacrimal fossa nfo narial fossa.
Figure 135.
Figure 135.
Jugal of Carcharodontosaurus saharicus (UCRC PV12). Left jugal in (A) lateral and (B) medial view. Right jugal in (C) lateral and (D) medial view. Scale bar equals 20 cm.
Figure 136.
Figure 136.
Postorbital of Carcharodontosaurus saharicus (UCRC PV12). Left postorbital in (A) lateral and (B) medial view. Right postorbital in (C) lateral and (D) medial view. Scale bar equals 20 cm. Abbreviations: aj articular surface for the jugal al articular surface for the lacrimal als articular surface for the lateralsphenoid asq articular surface for the squamosal ipr intraorbital process ob orbital brow vg vascular groove.
Figure 137.
Figure 137.
Posterior skull elements of Carcharodontosaurus saharicus (UCRC PV12). Braincase in (A) ventral, (B) anteroventral view and (C) posterior view. Scale bar equals 10 cm. Abbreviations: f frontal fm foramen magnum is interorbital septum popr paroccipital process.
Figure 138.
Figure 138.
Dentary fragments likely referable to Carcharodontosaurus saharicus. NMC 41859 in (A) right lateral, (B) medial and (C) dorsal (occlusal) view. FSAC-KK 02 in (D) dorsal (occlusal) and (E) medial view. NMC 41859 matched with the dentary of Giganotosaurus carolinii (MUCPv-95, redrawn and reversed from Calvo and Coria 1998) in (F) right lateral, (G) medial and (H) dorsal (occlusal) view. Scale bars equal 10 cm in A-E and 10 cm in F-H. F-G adjusted to same size (MUCPv-95 is 610 mm long, Calvo and Coria 1998).
Figure 139.
Figure 139.
Preliminary skeletal silhouette reconstruction of Carcharodontosaurus saharicus with preserved bones (including tentative referrals) shown in gray (after Ibrahim et al. 2017). Scale bar equals 1 m.
Figure 140.
Figure 140.
Reconstruction of the cranium of Carcharodontosaurus saharicus in left lateral view (premaxilla shortened, after Sereno et al. 1996).
See this image and copyright information in PMC

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