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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


Geographical setting of the Kem Kem region. Cenomanian (~94 Mya) paleogeographic world map showing key localities (map after


Outcrops of the Kem Kem sequence near Gara Sbaa. The red beds are overlain by a Cenomanian-Turonian limestone platform.


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.


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.


Examples of specimen measurements. A Turtle carapace fragment (MRS 172,


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.


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.


Landsat image of southeastern Morocco showing key localities, villages and cities, modified from


Jagged limestone talus from the overlying Cenomanian-Turonian platform at Gara Sbaa obscures the majority of the outcrop of the Kem Kem Group.


The escarpment at Gara Sbaa serves as a geographic landmark in the Kem Kem region.


Paleozoic strata in the Kem Kem and Tafilalt regions. A Unconformity at Iferda N’Ahouar B Devonian outcrop at Hmar Lakhdad (Fig.


Tectonic setting of the Kem Kem region. A Major tectonic complexes in southeastern Morocco (modified from


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.


Northern lithostratigraphic sections of the Kem Kem Group. Vertical scale bar equals 100 m. Abbreviations: c clay p pebble sd sand st silt.


Southern lithostratigraphic sections of the Kem Kem Group. Vertical scale bar equals 100 m. Abbreviations: c clay p pebble sd sand st silt.


Paleozoic rocks underlying the Kem Kem Group. A Jbel Sdila locality. B Close-up of Paleozoic basement.


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.


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.


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.


Tabular cross-bedding in the Gara Sbaa Formation. A Weathering of common tabular cross-beds B Close-up view of tabular cross-beds.


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.


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.


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.


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).


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.


Oum Tkout locality where quarrying has recovered an abundance of exceptionally preserved aquatic vertebrates and invertebrates in a still-water setting.


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.


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.


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.


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.


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 (


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.


Surface and in situ collecting. A Surface collecting at Gara Sbaa B Collecting in situ fossils at Boumerade.


Taphonomy at Oum Tkout. A Indeterminate paraclupeid fish with mineralized muscles (white) B Soft tissue (muscle) preservation under high magnification C Complete decapod (


Invertebrate boring structures in dinosaur bone. A Large bone piece showing local concentration of burrows (


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 (


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 (


Abrasion of Kem Kem fossils (FSAC-KK collection). A All localities B Gara Sbaa. X-axis is the abrasion index (0-4; see Table


Abrasion of Kem Kem fossils (FSAC-KK collection). A Boumerade. B Aferdou N’Chaft. X-axis is the abrasion index (0-4; see Table


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


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.


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.


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.


Possible biofilm (bacteria, fungi) from Oum Tkout, Douira Formation. Scale bar equals 5 mm.


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.


Decapod prawn Cretapenaeus berberus (


Dragonfly larva (Odonata indet.) from Oum Tkout, Douira Formation. Scale bar equals 5 mm.


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 (


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.


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.


Serenoichthys kemkemensis Dutheil, 1999 from the Douira Formation. Scale bar equals 1 cm.


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.


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.


Scales of a large holostean from the Kem Kem Group. A FSAC-KK 530 (Gara Sbaa) B NMC 50434-A C FSAC-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.


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 (


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.


Diplospondichthys moreaui Filleul and Dutheil, 2004 from the Kem Kem Group. Scale bar equals 2 cm.


Spinocaudichthys oumtkoutensis Filleul and Dutheil, 2002 from the Kem Kem Group. Scale bar equals 5 mm.


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


Large specimens from the Kem Kem Group likely referable to Axelrodichthys. Possible fragment of palatoquadrate region preserving a large part of the pterygoid (


Size variation in lungfish tooth plates from the Kem Kem Group. A


Amphibia and Squamata from the Douira Formation. Cf. Kababisha sp. braincase (


Testudinid shell fragments from the Kem Kem Group. Partial plastron with sutures (


Possible araripemydid and podocnemidinuran shell fragments from the Kem Kem Group. Possible araripemydid plastral element (


Isolated turtle postcrania from the Kem Kem Group. Pelomedusoides indet., cervical vertebra (


Partial shell and girdle bones of cf. Galianemys sp. (


Turtle skull elements from the Kem Kem Group. One of two undescribed skulls likely referable to Galianemys whitei (


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.


Sebecid skull, referred to Hamadasuchus rebouli (


Other sebecid cranial specimens. Posterolateral skull roof (FSAC-KK 930) in (A) dorsal and (B) right lateral view. Partial braincase of Hamadasuchus rebouli (


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.


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 (


Skull of Hamadasuchus rebouli (


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.


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.


Mandibular symphyses of Hamadasuchus rebouli. Mandibular symphysis (


Specimens of Laganosuchus from the Echkar Formation in Niger and the Kem Kem Group in Morocco. Cast of lower jaws (


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.


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.


Cranial remains of the crocodyliform Elosuchus. A, C E. cherifiensis cranium (


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 (


Braincase and jugal of the crocodyliform Elosuchus cherifiensis. Braincase (


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 (


Dentaries of the crocodyliform Elosuchus cherifiensis. A Left anterior dentary (


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.


Isolated pholidosaurid crocodyliform teeth. A NMC 41791 (see Fig.


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).


Ornithocheirid pterosaur teeth from the Kem Kem Group. A FSAC-KK 885 B FSAC-KK 44 C FSAC-KK 886 D, E FSAC-KK 197 in labial and lingual views F FSAC-KK 887 G FSAC-KK 941 H Teeth of ‘morphotype 1’ of


Ornithocheirid pterosaur jaw fragments from the Kem Kem Group. Premaxillae of Siroccopteryx moroccensis in (A) left lateral and (B) ventral views (from


Rostral fragment (BSP 1993 IX 338) referred by


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.


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.


Fragment, coated in ammonium chloride, of the mandibular symphysis of Xericeps curvirostris,


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.


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.


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.


Possible thyreophoran tooth.


Holotype material of Rebbachisaurus garasbae. A Dorsal vertebra (


Left humerus of Rebbachisaurus garasbae.


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 (


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.


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-S


Proximal section of a right ulna of a giant sauropod.


Indeterminate theropod teeth from the Kem Kem Group.


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 N MPDM 43 in lateral view O


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 (


Abelisaurid maxillae from Madagascar and Niger. A Left maxilla of Majungasaurus crenatissimus (


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.


Excavation and skeletal reconstruction of Deltadromeus agilis. A Excavation of the fibula B Mounted skeletal reconstruction using casts of the holotype


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.


Left metatarsals of Deltadromeus agilis (


Deltadromeus agilis from Morocco and Egypt. A Revised reconstruction based on


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.


Dentary fragments of Spinosaurus aegyptiacus from the Kem Kem Group. Dentary section (MPDM 30) in (A) lateral and (B) ?anterior view. Dentary fragment (


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.


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 (contra


Comparison of Spinosaurus aegyptiacus rostra from Morocco and Algeria. Moroccan rostra include


Rostral portions of Spinosaurus from Morocco and Algeria. Rostrum (


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.


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.


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 (


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.


Skull and skeleton of Spinosaurus aegyptiacus. A Skull reconstruction in left lateral view (after


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.


Teeth from Morocco and Niger referred to Carcharodontosaurus. Teeth of


Endocasts of Carcharodontosaurus saharicus. Endocast (from


Right maxilla of Carcharodontosaurus saharicus.


Nasal and lacrimal of Carcharodontosaurus saharicus (


Jugal of Carcharodontosaurus saharicus (


Postorbital of Carcharodontosaurus saharicus (


Posterior skull elements of Carcharodontosaurus saharicus (


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


Preliminary skeletal silhouette reconstruction of Carcharodontosaurus saharicus with preserved bones (including tentative referrals) shown in gray (after


Reconstruction of the cranium of Carcharodontosaurus saharicus in left lateral view (premaxilla shortened, after