Ouled Abdoun Basin

Ouled Abdoun Basin
Stratigraphic range: Maastrichtian-Ypresian
~
The Oulad Abdoun and other major phosphate basins (in yellow) of Morocco
TypeGeological formation
Unit ofSedimentary Basins
Lithology
PrimaryPhosphorite
OtherClay, Limestone, Sandstone
Location
Coordinates32°53′N 6°55′W / 32.883°N 6.917°W / 32.883; -6.917
Approximate paleocoordinates24°2′N 6°8′W / 24.033°N 6.133°W / 24.033; -6.133
RegionBéni Mellal-Khénifra, Khouribga
Country Morocco
Ouled Abdoun Basin (Morocco)

The Oulad Abdoun Basin (also known as the Ouled Abdoun Basin or Khouribga Basin) is a phosphate sedimentary basin located in Morocco, near the city of Khouribga. It is the largest in Morocco, comprising 44% of Morocco's phosphate reserves, and at least 26.8 billion tons of phosphate.[1][2] It is also known as an important site for vertebrate fossils, with deposits ranging from the Late Cretaceous (Maastrichtian) to the Eocene epoch (Ypresian), a period of about 25 million years.[3]

Geography

The Oulad Abdoun is located west of the Atlas Mountains, near the city of Khouribga. The Oulad Abdoun phosphate deposits encompass some 100 by 45 kilometres (62 by 28 mi), an area of 4,500 square kilometres (1,700 sq mi).[2] The Oulad Abdoun is the largest and northernmost of Morocco's major phosphate basins, which from northeast to southwest, include the Ganntour, Meskala, and Oued Eddahab (Laayoune-Baa) basins.[1][2]

Paleobiota

Life restoration of Ouled Abdoun Basin paleoenvironments during the late Cretaceous: dinosaurs of Sidi Chennae (left) and numerous aquatic animals of Sidi Daoui (right).
Color key
Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
Notes
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

The Oulad Abdoun Basin stretches from the late Cretaceous to the Eocene and contains abundant marine vertebrate fossils, including sharks, bony fish, turtles, crocodilians, and other reptiles, as well as sea birds and a small number of terrestrial mammals.[3][4]

Molluscs

Mollusca from the Khouribga Phosphates[5]
Genus Species Location Time Material Notes Images
Cucullaea (Idonearca) C. (I.) thevestensis A false ark shell.
Heligmopsis H. wegmanianus Maastrichtian A feather oyster also present in Ganntour basin.
Venericardia V. coquandi Danian A carditid clam also present in Ganntour basin.
Baculites B. sp. Maastrichtian A straight-shelled ammonite, also present in Ganntour basin.
Cimomia C. aff. sudanensis Ypresian A nautiloid also present in Ganntour basin.
Ampullinopsis A. crassatina An ampullinid gastropod.
Hemithersitea H. chouberti Lutetian A thersiteid gastropod.

Fish

Bony fish

Bony fish from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Acipenseridae[6] Indeterminate Maastrichtian First record of a sturgeon on the continent of Africa.
Enchodus E. libycus Maastrichtian An enchodontid aulopiform.
E. bursauxi Maastrichtian
E. elegans Maastrichtian
Eoserrasalmimus[7] E. cattoi Maastrichtian A pycnodontiform.
Mawsoniidae[8] Indeterminate Maastrichtian A giant mawsoniid coelacanth measuring 3.65–5.52 metres (12.0–18.1 ft) long.
Phacodus[9] P. punctatus var. africanus Maastrichtian A pycnodontiform.
Plethodidae[10] Indeterminate Maastrichtian Youngest occurrence of plethodid fish.
Pycnodontidae Indeterminate Maastrichtian
Pseudoegertonia P. sp. Maastrichtian
Serrasalmimus[7] S. secans Thanetian A carnivorous pycnodontiform.
Stephanodus S. libycus Maastrichtian A pycnodontid
Stratodus S. apicalis Maastrichtian An alepisauroform.
Macroprosopon M. hiltoni[11] Ypresian A marine osteoglossid.

Sharks

Sharks from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Abdounia[12] A. africana Selandian A requiem shark.
Brachycarcharias B. atlasi[11] Ypresian
Casieria C. maghrebiana Selandian
Chiloscyllium C. salvani. Selandian A carpet shark.
Cretalamna C. appendiculata Maastrichtian An otodontid shark
C. maroccana Maastrichtian
Delpitoscyllium D. africanum Selandian
Eostegostoma E. sp. Selandian A carpet shark.
Foumtizia F. abdouni Selandian
Galeorhinus G. mesetaensis Selandian A houndshark
Ginglymostoma G. chenanei Selandian An early relative of nurse shark
G. khouribgaense Selandian
Hologinglymostoma H. jaegeri Selandian
Khouribgaleus K. gomphorhiza Selandian
Metlaouia M. delpiti Selandian
Microscyliorhinus M. simplex Selandian
Palaeogaleus P. larachei Selandian
Palaeorhincodon P. daouii Selandian A carpet shark.
Porodermoides P. spanios Selandian
Premontreia P. peypouqueti Selandian
P. subulidens Selandian
Pseudocorax[13] P. heteroserratus Maastrichtian A pseudocoracid mackerel shark
Scyliorhinus S. ptychtus Selandian A catshark
S. entomodon Selandian
S. sulcidens Selandian
Serratolamna S. serrata Maastrichtian A mackerel shark
S. khderii Maastrichtian
Squalicorax S. pristodontus Maastrichtian A crow shark
S. bassanii Maastrichtian
S. africanus Maastrichtian
S. microserratus Maastrichtian
S. benguerirensis Maastrichtian
Triakis T. antunesi Selandian A houndshark

Other cartilaginous fish

Other cartilaginous fish from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Archaemanta A. priemi Selandian A ray.
Burnhamia B. cf. davisi Selandian A devil ray
Coupatezia C. larivei Selandian A ray.
C. fallax Maastrichtian
C. elevata Maastrichtian
Dasyatis D. ponsi Selandian A stingray. An indeterminate species found in Maastrichtian deposits of Ganntour Basin.
Delpitia D. reticulata Selandian A ray.
Gymnura G. delpiti Selandian A ray.
Heterobatis H. talbaouii Selandian A ray.
Heterotorpedo H. brahimi Selandian A ray.
Hypolophodon H. sp. Selandian A ray.
Rhombodus R. binkhorsti Maastrichtian A ray.
R. microdus Maastrichtian
R. meridionalis Maastrichtian
Schizorhiza S. stromeri Maastrichtian A sclerorhynchoid skate which resembled a sawfish.

Dinosaurs

Ornithischians

Ornithischians from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Ajnabia[14] A. odysseus
  • Office Cherifien des Phosphates mines at Sidi Chennane
 Maastrichtian
  • partial left maxilla with teeth
  • fragments of right maxilla
  • partial right dentary
 A small lambeosaurine hadrosaur; the first known from Africa
Lambeosaurinae[15] Indeterminate
  • Mrah Lahrach
 Maastrichtian
  • right femur
 A lambeosaurine larger than Ajnabia and Minqaria
Lambeosaurinae[15] Indeterminate
  • Sidi Daoui
 Maastrichtian
  • left humerus
 A lambeosaurine larger than Ajnabia and Minqaria
Minqaria[15] M. bata
  • Sidi Chennane
 Maastrichtian
  • right maxilla
  • left dentary
  • braincase
 A lambeosaurine hadrosaur
Taleta[16] T. taleta
  • Sidi Chennane
 Maastrichtian
  • partial left and right maxilla with teeth
 A lambeosaurine hadrosaur

Sauropods

Sauropods from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Titanosauria[17] Indeterminate Maastrichtian
  • right femur, tibia, and fibula
  • undescribed material
 A new titanosaurian taxon, currently unnamed.

Theropods

Abelisaurs
Abelisaurs from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Abelisauridae[19] Indeterminate
  • Sidi Chennane
 Maastrichtian right tibia Probably distinct from Chenanisaurus due to its smaller size (about 5 m (16 ft) long) despite being mature. Similar to South American abelisaurids. Possibly a member of Furileusauria.[20]
Abelisauridae[19] Indeterminate
  • Sidi Daoui
 Maastrichtian right metatarsal II A mature specimen but very small for an abelisaurid, about 2.6 m (8.5 ft) long
Chenanisaurus[17] C. barbaricus
  • Sidi Chennane
  • Sidi Daoui
 Maastrichtian
  • anterior end of left dentary
  • 2 premaxillary teeth
  • a maxillary tooth
 A large abelisaurid theropod.
Birds

Bird fossils are common in the Basin, which includes the oldest birds in Africa.[4] At least three orders and several families of sea birds are represented, including Procellariiformes (albatrosses and petrels, fossils assignable to Diomedeidae and Procellariidae), Pelecaniformes (pelicans and allies, fossils assignable to Phaethontidae, Prophaethontidae, Fregatidae and Pelagornithidae), and Anseriformes (waterfowl, including fossil Presbyornithidae).[4]

Birds from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Dasornis[21] D. abdoun Sidi Daoui Ypresian A holotype specimens consists of carpometacarpus, ulna, radius, tibiotarsus, and humerus. A pelagornithid, or pseudotooth bird.
D. emuinus Sidi Daoui Ypresian A partial skeleton with incomplete cranium.
D. tolapica Selandian A parital skeleton.
Lithoptila[22] L. abdounensis Sidi Daoui Thanetian A neurocranium. A prophaethontid seabird related to modern tropicbirds.

Pterosaurs

Pterosaurs from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Alcione A. elainus[23] Sidi Daoui Maastrichtian A nyctosaurid which shows possible adaptations towards diving behaviour.
Barbaridactylus B. grandis[23] Sidi Daoui Maastrichtian A nyctosaurid.
Simurghia S. robusta[23] Sidi Daoui Maastrichtian A nyctosaurid related to Alcione.
Phosphatodraco P. mauritanicus[24] Sidi Daoui Maastrichtian An azhdarchid.
cf Arambourgiania[23] A. sp? Sidi Chennane Maastrichtian A giant azhdarchid, may belong to the Jordanian taxon Arambourgiania.
aff. Quetzalcoatlus[23] indet. Maastrichtian
Tethydraco T. regalis[23] Sidi Daoui Maastrichtian A pterosaur of debated affinity; may be an azhdarchid[25] or a pteranodontid.[23]

Crocodylomorphs

Crocodylomorphs from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Arambourgisuchus[26] A. khouribgaensis Thanetian A dyrosaurid.
Atlantosuchus A. coupatezi Danian A dyrosaurid.
Chenanisuchus C. lateroculi Thanetian The most basal known dyrosaurid.
Dyrosaurus D. maghribensis Ypresian A dyrosaur.
Maroccosuchus[27] M. zennaroi Ypresian A tomistomine crocodylian.
Ocepesuchus[28] O. eoafricanus Maastrichtian A gavialoid crocodylian.

Squamates

Mosasaurs

The late Maastrichtian deposits of the Khouribga Phosphates preserve abundant mosasaur fossils, especially isolated teeth. At least fifteen species are represented, covering the majority of ecological niches the group is known to have occupied.[29] The Ouled Abdoun Basin is one of the richest known mosasaur fossil sites in the world.[29]

Mosasaurs from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Carinodens[30][29] C. acrodon Maastrichtian A small mosasaurine mosasaur with crushing teeth.[31] Fossils of all reported Carinodens species are very rare in Khouribga.[31] The dentition differs slightly between species, suggesting marginally different diets.[31] More fossil material is required to determine if both C. minalmamar and C. belgicus are present in the Khouribga Phosphates, as the present material is difficult to differentiate.[29]
C. belgicus
C. minalmamar
Eremiasaurus E. heterodontus Maastrichtian
  • Partial and subcomplete skeletons[31]
  • Isolated teeth[31]
 A mosasaurine mosasaur with unusually heterodont teeth, including straight and conical anterior teeth, blade-like median teeth, and recurved posterior teeth.[31] A medium-sized (4.5–5 metres) predator and one of the more commonly found fossil mosasaurs in the Khouribga Phosphates.[31]
Gavialimimus[32] G. almaghribensis Maastrichtian An unusual longirostrine plioplatecarpine mosasaur, measuring about 6 metres long.[31] Fossils of G. almaghribensis are very common in the Khouribga Phosphates.[31]
Globidens G. phosphaticus Maastrichtian A durophagous mosasaurine mosasaur with specialized crushing teeth.[31] G. phosphaticus is known from Morocco only in the form of isolated teeth but more complete material from the species is known from Angola.[31] G. phosphaticus and G. simplex were of similar size but had slightly different teeth, suggesting marginally different diets.[31]
G. simplex Maastrichtian
Halisaurus H. arambourgi Maastrichtian
  • Several skeletons[31]
  • Isolated teeth[31]
 A halisaurine mosasaur, measuring about 3–4 metres long.[31] One of the most commonly found mosasaurs in Morocco.[31]
Igdamanosaurus I. aegyptiacus Maastrichtian "Globidens aegyptiacus" was previously reported as present in the Khouribga Phosphates on the basis of isolated teeth. G. aegyptiacus was reclassified as Igdamanosaurus aegyptiacus in 1991 and differs from the globidensin material known from Morocco.[33] Moroccan fossils previously referred to G. aegyptiacus are now referred to G. phosphaticus.[33]
Khinjaria[34] K. acuta
  • Sidi Chennane
 Maastrichtian
  • A partial skull and vertebra[34]
 A plioplatecarpine mosasaur, known only from a single specimen.[31] Khinjaria possessed a unique skull and teeth morphology and might have been an ambush predator.[31]
Liodon L. anceps Maastrichtian "Mosasaurus (Leiodon) cf. anceps" was reported as present in Khouribga by Camille Arambourg (1952) based on isolated teeth.[35] Liodon anceps is now considered a nomen dubium.[31] Teeth previously referred to Liodon likely belong to Eremiasaurus heterodontus (small and slender specimens) and Thalassotitan atrox (other specimens).[31]
Mosasaurus M. beaugei Maastrichtian
  • Skulls and mandibles[31]
  • Isolated teeth[31]
 A large mosasaurine mosasaur, measuring about 8–10 metres long.[31] Relatively scarce compared to other mosasaurs of comparable size. Likely a local apex predator.[31]
Platecarpus P. ptychodon Maastrichtian
  • Isolated teeth and vertebrae[31]
 Platecarpus ptychodon is known only from isolated material. The fossils referred to P. ptychodon are treated either as non-diagnostic (in which case P. ptychodon is a nomen dubium) or as belonging to Gavialimimus almaghribensis (in which case G. almaghribensis is considered a junior synonym of the new combination Gavialimimus ptychodon).[31]
Pluridens[36] P. serpentis Maastrichtian
  • Complete skulls with associated vertebrae[31]
  • Isolated teeth[31]
 A large halisaurine mosasaur, also known from Niger and Nigeria. Measured about 5–6 metres long.[31] Pluridens is relatively uncommon in Morocco and distinguished from the closely related Halisaurus based on features in the skull and its larger size.[31] The teeth of Pluridens are identical to those of Halisaurus, only larger, and the two are thus difficult to differentiate in samples of isolated teeth.[31]
P. imelaki[37]
  • Skull and mandibles
Prognathodon P. currii Maastrichtian A large mosasaurine mosasaur. Prognathodon curii is known in Morocco only from isolated teeth and is comparably rare, with only about a dozen teeth identified.[31] The related species Prognathodon giganteus is also known from Morocco, though only from earlier lower Maastrichtian deposits in the Gantour Basin to the southwest of the Ouled Abdoun Basin.[31][35]
Stelladens[38] S. mysteriosus Maastrichtian
  • A dentary fragment[31]
  • Isolated teeth[31]
 A medium-sized mosasaurine with distinct, unique teeth structure with prominent serrated ridges.[31] Stelladens is a very scarce taxon in the Khouribga Phosphates.[31] Stelladens would have measured about 5 metres long and was probably a highly specialized predator.[31]
Thalassotitan[39] T. atrox Maastrichtian
  • Several skeletons[31]
  • Isolated teeth[31]
 A large-bodied mosasaurine with powerful jaws and massive teeth, closely related to Prognathodon.[31] Fossils of Thalassotitan are comparably very common and it likely represented the largest local apex predator.[31] Fossils now referred to Thalassotitan were before its description often referred to as "Prognathodon nov. sp."[31]
Xenodens[40] X. calminechari Maastrichtian
  • A left maxilla with four teeth[31]
  • A left maxilla with eight teeth[41]
 A small mosasaurine mosasaur with unusual, shark-like teeth.[31] The holotype of Xenodons was suggested to potentially be a forged specimen by Sharpe et al. (2024),[42] but Longrich et al. (2025) published a rebuttal based on CT scans of the holotype and another referred specimen, arguing for the validity of the genus.[41]

Snakes

Snakes from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Palaeophis[43] P. maghrebianus Maastrichtian?–Ypresian Multiple fossilized vertebrae. A palaeophiid marine snake.

Varanoids

Varanoids from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Pachyvaranus P. crassispondylus Maastrichtian
  • Vertebrae and ribs[44]
 A near-shore marine lizard.[44] Originally described based only on a small number of isolated vertebrae. Discovery of further vertebrae and ribs in Morocco and Syria allowed in 2011 for classification in the Varanoidea, in a new family Pachyvaranidae.[44] Osteoderms were originally referred to this taxon but likely come from fish instead, perhaps Stratodus.[44]

Plesiosaurs

Plesiosaur material found in the Ouled Abdoun Basin has been referred to one valid species, Zarafasaura oceanis. Isolated elasmosaurid fossils may all belong to this species but comparison is difficult.[45][46]

Plesiosaurs from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Elasmosauridae indet. Sidi Daoui Maastrichtian
  • Articulated and isolated vertebrae[46]
  • Pelvic elements[46]
 Elasmosaurid fossils without cranial elements. Could represent the same taxon as Zarafasaura oceanis but comparison impossible at the time of discovery due to no overlap in the fossil material.[46] Comparison may be possible after the discovery of postcranial Zarafasaura elements in 2013.[45]
Plesiosaurus P. mauritanicus Maastrichtian
  • Isolated teeth and vertebrae[45]
 Named by Camille Arambourg (1952) based on isolated fossils. Plesiosaurus mauritanicus is regarded as a nomen dubium since the fossils are non-diagnostic beyond being elasmosaurid, and because the holotype and paratypes are from different individuals found at different localities.[45] Potentially the same taxon as Zarafasaura oceanis.[45]
Zarafasaura Z. oceanis Sidi Daoui Maastrichtian
  • Holotype specimen, a crushed skull and partial mandible[45]
  • Paratype specimen, complete skull with many postcranial elements[45]
 A relatively small elasmosaurid plesiosaur, closely related to plesiosaurs known from North America and Japan.[47]

Turtles

Turtles from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Alienochelys[48] A. selloumi Sidi Daoui Maastrichtian A sea turtle with crushing jaws
Araiochelys[49] A. hirayamai Sidi Daoui Maastrichtian A skull, lower jaws, partial disarticulated shell, including a few carapace fragments (left first and fifth costals, left third, seventh, and eighth peripherals and other fragments), nearly complete plastron, and limb bones. A bothremydid pleurodiran turtle
Argillochelys[50] A. africana Ypresian A pancheloniid sea turtle
Bothremys[49] B. kellyi Danian-Thanetian A bothremydid pleurodiran turtle
B. maghrebiana
Brachyopsemys B. tingitana Danian A sandownid sea turtle
Euclastes E. acutirostris Danian-Thanetian A pancheloniid sea turtle. An indeterminate species also present in the Maastrichtian deposits of Ganntour Basin.[51]
E. wielandi Danian
Labrostochelys[49] L. galkini Oued Zem Danian A nearly complete skull A bothremydid pleurodiran turtle
Marocokatognathus[52] M. jimenezi near Oued Zem Thanetian–Ypresian A partial mandible A cheloniid sea turtle
Ocepechelon[53] O. bouyai Sidi Chennane Maastrichtian A complete but isolated 70-cm-long skull. A giant protostegid sea turtle with an unusual pipette-like snout
Puppigerus P. camperi Ypresian A Pancheloniid sea turtle
Rhothonemys[49] R. brinkmani Danian A bothremydid pleurodiran turtle
Tasbacka T. ouledabdounensis Danian A cheloniid sea turtle
Taphrosphys T. ippolitoi Danian A bothremydid pleurodiran turtle
Ummulisani U. rutgersensis Mrah Iaresh Ypresian A skull without a palate A bothremydid pleurodiran turtle

Mammals

Afrotheres

Afrotheres from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Abdounodus[54] A. hamdii Thanetian-Ypresian A lower jaw. An early afrotherian mammal.
Daouitherium[55] D. rebouli Ypresian lower jaws and associated cheek teeth An early proboscidean.
Eritherium[56] E. azzouzorum Sidi Chennane Thanetian An upper jaw (with approaches of the zygomatic bone and two maxillary branches, each of the two posterior premolars (P3 and 4) and three molars (M1-3)) An early proboscidean.
Hadrogeneios[57] H. phosphaticus Selandian-Thanetian A mandibular symphysis with some partial left dentary, two sockets for the incisors, the first lower canine, and the first to third premolars An early afrotherian mammal.
Ocepeia[54] O. daouiensis Grand Daoui, Meraa El Arech, and Sidi Chennane Selandian Partial skull consists of an upper jaw and dental fragments An early afrotherian mammal.

O. grandis Thanetian Lower jaw and teeth.
Phosphatherium P. escuilliei Selandian An early proboscidean.
Stylolophus[58] S. minor Ypresian An embrithopod.
S. major Ypresian

Hyaenodonts

Hyaenodonts from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Boualitomus B. marocanensis Selandian A hyaenodont.
Lahimia[59] L. selloumi Selandian A hyaenodont.

See also

References

  1. ^ a b S. Zouhri; A. Kchikach; O. Saddiqi; F.Z. El Haimer; L. Baidder; A. Michard (2008). "The Cretaceous-Tertiary Plateaus". In A. Michard; O. Saddiqi; A. Chalouan; D. Frizon de Lamotte (eds.). Continental Evolution: the Geology of Morocco. Berlin: Springer. pp. 331–358. ISBN 978-3-540-77075-6.
  2. ^ a b c Office Chérifien des Phosphates (1989). "The Phosphate Basins of Morocco". In A.J.G. Notholt; R.P. Sheldon; D.F. Davidson (eds.). Phosphate Deposits of the World. Volume 2, Phosphate Rock Resources. Cambridge, Eng.: Cambridge University Press. pp. 301–311. ISBN 978-0-521-67333-4.
  3. ^ a b Yans, Johan; Amaghzaz, M'Barek; Bouya, Baadi; Cappetta, Henri; Iacumin, Paola; Kocsis, László; Mouflih, Mustapha; Selloum, Omar; Sen, Sevket; Storme, Jean-Yves; Gheerbrant, Emmanuel (2014). "First carbon isotope chemostratigraphy of the Ouled Abdoun phosphate Basin, Morocco; implications for dating and evolution of earliest African placental mammals". Gondwana Research. 25 (1): 257–269. Bibcode:2014GondR..25..257Y. doi:10.1016/j.gr.2013.04.004.
  4. ^ a b c Gheerbrant, E.; Sudre, J.; Cappetta, H.; Mourer-Chauviré, C.; Bourdon, E.; Iarochene, M.; Amaghzaz, M.; Bouya, B. (2003). "Les localités à mammifères des carrières de Grand Daoui, bassin des Ouled Abdoun, Maroc, Yprésien: premier état des lieux" [The mammal localities of Grand Daoui Quarries, Ouled Abdoun Basin, Morocco, Ypresian: A first survey] (PDF). Bull. Soc. Géol. Fr. (in French and English). 174 (3): 279–293. doi:10.2113/174.3.279.
  5. ^ Pacaud, J.M.; Lebrun, P. (2020). "Moroccan phosphate beds: hot-spot for Maastrichtian-Eocene fossils. The Invertebrate fauna.". In Lebrun, P. (ed.). Fossils from Morocco, Volume IIa, Emblematic localities from the Mesozoic and the Paleogene. Les Éditions du Piat. pp. 235–245. ISBN 978-2-917198-46-9.
  6. ^ Martill, D. M. (2023). "A sturgeon (Actinopterygii, Acipenseriformes) from the Upper Cretaceous of Africa". Cretaceous Research. 148 105546. 105546. Bibcode:2023CrRes.14805546M. doi:10.1016/j.cretres.2023.105546. S2CID 257863907.
  7. ^ a b Vullo, Romain; Cavin, Lionel; Khalloufi, Bouziane; Amaghzaz, Mbarek; Bardet, Nathalie; Jalil, Nour-Eddine; Jourani, Essaid; Khaldoune, Fatima; Gheerbrant, Emmanuel (2017-07-28). "A unique Cretaceous–Paleogene lineage of piranha-jawed pycnodont fishes". Scientific Reports. 7 (1): 6802. Bibcode:2017NatSR...7.6802V. doi:10.1038/s41598-017-06792-x. ISSN 2045-2322. PMC 5533729. PMID 28754956.
  8. ^ Brito, P.M.; Martill, D.M.; Eaves, I.; Smith, R.E.; Cooper, S.L.A. (2021). "A marine Late Cretaceous (Maastrichtian) coelacanth from North Africa". Cretaceous Research. 122 104768. Bibcode:2021CrRes.12204768B. doi:10.1016/j.cretres.2021.104768. S2CID 233551515.
  9. ^ Cooper, Samuel L. A.; Marson, Kyle J.; Smith, Roy E.; Martill, David (2022-03-01). "Contrasting preservation in pycnodont fishes reveals first record of regurgitalites from the Upper Cretaceous (Maastrichtian) Moroccan phosphate deposits". Cretaceous Research. 131 105111. Bibcode:2022CrRes.13105111C. doi:10.1016/j.cretres.2021.105111. ISSN 0195-6671.
  10. ^ Cooper, Samuel. L. A.; Norton, Jack L. (2023-08-08). "Youngest occurrence of a plethodid fish (Teleostei: Tselfatiiformes: Plethodidae) from the Maastrichtian of North Africa". Cretaceous Research. 152 105673. Bibcode:2023CrRes.15205673C. doi:10.1016/j.cretres.2023.105673. ISSN 0195-6671.
  11. ^ a b Capobianco, A.; Zouhri, S.; Friedman, M. (2024). "A long-snouted marine bonytongue (Teleostei: Osteoglossidae) from the early Eocene of Morocco and the phylogenetic affinities of marine osteoglossids". Zoological Journal of the Linnean Society. 203 zlae015. doi:10.1093/zoolinnean/zlae015.
  12. ^ Noubhani, A.; Cappetta, H. (1997). "Les Orectolobiformes, Carcharhiniformes et Myliobatiformes (Elasmobranchii, Neoselachii) des bassins à phosphate du Maroc (Maastrichtien-Lutétien basal). Systématique, biostratigraphie, évolution et dynamique des faunes". Palaeo Ichthyologica.
  13. ^ Egli, H. C.; Goode, B.; Rempert, T.; Rego, C. (2025). "Pseudocorax (Chondrichthyes, Lamniformes, Pseudocoracidae) in the Upper Maastrichtian phosphates of Khouribga Province, Morocco". Journal of Paleontology: 1–15. doi:10.1017/jpa.2025.10183.
  14. ^ Longrich, Nicholas R.; Suberbiola, Xabier Pereda; Pyron, R. Alexander; Jalil, Nour-Eddine (2020). "The first duckbill dinosaur (Hadrosauridae: Lambeosaurinae) from Africa and the role of oceanic dispersal in dinosaur biogeography". Cretaceous Research. 120 104678. doi:10.1016/j.cretres.2020.104678. S2CID 228807024.
  15. ^ a b c Longrich, N. R.; Pereda-Suberbiola, X.; Bardet, N.; Jalil, N.-E. (2024). "A new small duckbilled dinosaur (Hadrosauridae: Lambeosaurinae) from Morocco and dinosaur diversity in the late Maastrichtian of North Africa". Scientific Reports. 14 (1) 3665. 3665. Bibcode:2024NatSR..14.3665L. doi:10.1038/s41598-024-53447-9. PMC 10864364. PMID 38351204.
  16. ^ Longrich, Nicholas R.; Pereda-Suberbiola, Xabier; Bardet, Nathalie; Jalil, Nour-Eddine (2025-05-28). "A new hadrosaurid dinosaur from the late Maastrichtian Phosphates of Morocco provides evidence for an African radiation of lambeosaurines". Gondwana Research. 145 (in press): 142–151. Bibcode:2025GondR.145..142L. doi:10.1016/j.gr.2025.05.006.
  17. ^ a b Longrich, N.R.; Pereda-Suberbiola, X.; Jalil, N.-E.; Khaldoune, F.; Jourani, E. (2017). "An abelisaurid from the latest Cretaceous (late Maastrichtian) of Morocco, North Africa". Cretaceous Research. 76: 40–52. Bibcode:2017CrRes..76...40L. doi:10.1016/j.cretres.2017.03.021.
  18. ^ Pereda Suberbiola, Xabier; Bardet, Nathalie; Iarochène, Mohamed; Bouya, Baâdi; Amaghzaz, Mbarek (2004-09-01). "The first record of a sauropod dinosaur from the Late Cretaceous phosphates of Morocco". Journal of African Earth Sciences. 40 (1): 81–88. Bibcode:2004JAfES..40...81P. doi:10.1016/j.jafrearsci.2004.07.002. ISSN 1464-343X.
  19. ^ a b Longrich, Nicholas R.; Isasmendi, Erik; Pereda-Suberbiola, Xabier; Jalil, Nour-Eddine (August 2023). "New fossils of Abelisauridae (Dinosauria: Theropoda) from the upper Maastrichtian of Morocco, North Africa". Cretaceous Research. 152 105677. Bibcode:2023CrRes.15205677L. doi:10.1016/j.cretres.2023.105677.
  20. ^ Buffetaut, E.; Tong, H.; Girard, J.; Hoyez, B.; Párraga, J. (2024). "Caletodraco cottardi: A New Furileusaurian Abelisaurid (Dinosauria: Theropoda) from the Cenomanian Chalk of Normandy (North-Western France)". Fossil Studies. 2 (3): 177–195. doi:10.3390/fossils2030009.
  21. ^ Bourdon, Estelle & Amaghzaz, M'barek & Bouya, Baâdi. (2010). Pseudotoothed Birds (Aves, Odontopterygiformes) from the Early Tertiary of Morocco. American Museum Novitates. 3704. 1-71. 10.1206/3704.2.
  22. ^ Bourdon, Estelle; Bouya, Baâdi; Iarochene, Mohamed (2005). "Earliest African neornithine bird: A new species of prophaethontidae (Aves) from the Paleocene of Morocco". Journal of Vertebrate Paleontology. 25: 157. doi:10.1671/0272-4634(2005)025[0157:EANBAN]2.0.CO;2. ISSN 0272-4634. S2CID 86218884.
  23. ^ a b c d e f g Longrich, Nicholas R.; Martill, David M.; Andres, Brian (2018). "Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary". PLOS Biology. 16 (3) e2001663. doi:10.1371/journal.pbio.2001663. PMC 5849296. PMID 29534059.
  24. ^ Suberbiola, Xabier Pereda; Bardet, Nathalie; Jouve, Stéphane; Iarochène, Mohamed; Bouya, Baâdi; Amaghzaz, Mbarek (2003). "A new azhdarchid pterosaur from the Late Cretaceous phosphates of Morocco". Geological Society, London, Special Publications. 217 (1): 79–90. Bibcode:2003GSLSP.217...79S. doi:10.1144/GSL.SP.2003.217.01.08. S2CID 135043714.
  25. ^ Labita, Claudio; Martill, David M. (October 2020). "An articulated pterosaur wing from the Upper Cretaceous (Maastrichtian) phosphates of Morocco". Cretaceous Research. 119 104679. doi:10.1016/j.cretres.2020.104679. S2CID 226328607.
  26. ^ Jouve, S.; Iarochène, M.; Bouya, B.; Amaghzaz, M. (2005). "A new dyrosaurid crocodyliform from the Palaeocene of Morocco and a phylogenetic analysis of Dyrosauridae". Acta Palaeontologica Polonica. 50 (3): 581–594.
  27. ^ Jonet, S.; Wouters, G. (1977). "Maroccosuchus zennaroi, crocodilien eusuchien nouveau des phosphates du Maroc". Notes et Mémoires du Service Géologique du Maroc. 38: 177–202.
  28. ^ Jouve, Stéphane; Bardet, Nathalie; Jalil, Nour-Eddine; Suberbiola, Xabier Pereda; Bouya, Baâdi; Amaghzaz, Mbarek (2008). "The oldest African crocodylian: phylogeny, paleobiogeography, and differential survivorship of marine reptiles through the Cretaceous-Tertiary boundary" (PDF). Journal of Vertebrate Paleontology. 28 (2): 409–421. doi:10.1671/0272-4634(2008)28[409:TOACPP]2.0.CO;2. S2CID 86503283.
  29. ^ a b c d e f Longrich, N. R.; Pereda-Suberbiola, X.; Jalil, N.-E.; Bardet, N. (2024). "A New Species of the Durophagous Mosasaurid Carinodens from the Late Maastrichtian Phosphates of Morocco and Implications for Maastrichtian Mosasaurid Diversity". Diversity. 17 (1). 25. Bibcode:2024Diver..17...25L. doi:10.3390/d17010025.
  30. ^ A.S. Schulp; N. Bardet; B. Bouya (2009). "A new species of the durophagous mosasaur Carinodens (Squamata, Mosasauridae) and additional material of Carinodens belgicus from the Maastrichtian phosphates of Morocco". Netherlands Journal of Geosciences. 88 (3): 161–167. Bibcode:2009NJGeo..88..161S. doi:10.1017/S0016774600000871. S2CID 130121783.
  31. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc Bardet, Nathalie; Fischer, Valentin; Jalil, Nour-Eddine; Khaldoune, Fatima; Yazami, Oussama Khadiri; Pereda-Suberbiola, Xabier; Longrich, Nicholas (2025). "Mosasaurids Bare the Teeth: An Extraordinary Ecological Disparity in the Phosphates of Morocco Just Prior to the K/Pg Crisis". Diversity. 17 (2): 114. Bibcode:2025Diver..17..114B. doi:10.3390/d17020114. ISSN 1424-2818.
  32. ^ Strong, Catherine R. C.; Caldwell, Michael W.; Konishi, Takuya; Palci, Alessandro (2020-09-28). "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". Journal of Systematic Palaeontology. 18 (21): 1769–1804. Bibcode:2020JSPal..18.1769S. doi:10.1080/14772019.2020.1818322. ISSN 1477-2019.
  33. ^ a b Bardet, N.; Suberbiola, X. Pereda; Iarochène, M.; Amalik, M.; Bouya, B. (2005). "Durophagous Mosasauridae (Squamata) from the Upper Cretaceous phosphates of Morocco, with description of a new species of Globidens". Netherlands Journal of Geosciences. 84 (3): 167–175. Bibcode:2005NJGeo..84..167B. doi:10.1017/S0016774600020953. ISSN 0016-7746.
  34. ^ a b Longrich, Nicholas R.; Polcyn, Michael J.; Jalil, Nour-Eddine; Pereda-Suberbiola, Xabier; Bardet, Nathalie (2024-03-01). "A bizarre new plioplatecarpine mosasaurid from the Maastrichtian of Morocco". Cretaceous Research. 160 105870. Bibcode:2024CrRes.16005870L. doi:10.1016/j.cretres.2024.105870. hdl:1874/438706. ISSN 0195-6671.
  35. ^ a b Bardet, Nathalie; Houssaye, Alexandra; Vincent, Peggy; Pereda Suberbiola, Xabier; Amaghzaz, Mbarek; Jourani, Essaid; Meslouh, Saïd (2015). "Mosasaurids (Squamata) from the Maastrichtian Phosphates of Morocco: Biodiversity, palaeobiogeography and palaeoecology based on tooth morphoguilds". Gondwana Research. 27 (3): 1068–1078. Bibcode:2015GondR..27.1068B. doi:10.1016/j.gr.2014.08.014. ISSN 1342-937X.
  36. ^ Longrich, N. R.; Bardet, N.; Khaldoune, F.; Khadiri Yazami, O.; Jalil, N.-E. (2021). "Pluridens serpentis, a new mosasaurid (Mosasauridae: Halisaurinae) from the Maastrichtian of Morocco and implications for mosasaur diversity". Cretaceous Research. 126 104882. Bibcode:2021CrRes.12604882L. doi:10.1016/j.cretres.2021.104882.
  37. ^ Longrich, N.R.; Jalil, N.-E. (2026). "A Giant Halisaurine from the Late Maastrichtian of Morocco". Diversity. 18 (3) 3. doi:10.3390/d18030159.
  38. ^ Longrich, Nicholas R.; Jalil, Nour-Eddine; Pereda-Suberbiola, Xabier; Bardet, Nathalie (2023). "Stelladens mysteriosus: A Strange New Mosasaurid (Squamata) from the Maastrichtian (Late Cretaceous) of Morocco". Fossils. 1 (1): 2–14. doi:10.3390/fossils1010002. hdl:10810/64193. ISSN 2813-6284.
  39. ^ Longrich, Nicholas R.; Jalil, Nour-Eddine; Khaldoune, Fatima; Yazami, Oussama Khadiri; Pereda-Suberbiola, Xabier; Bardet, Nathalie (24 August 2022). "Thalassotitan atrox, a giant predatory mosasaurid (Squamata) from the Upper Maastrichtian Phosphates of Morocco". Cretaceous Research. 140 105315. Bibcode:2022CrRes.14005315L. doi:10.1016/j.cretres.2022.105315. S2CID 251821884.
  40. ^ Longrich, N. R.; Bardet, N.; Schulp, A. S.; Jalil, N.-E. (2021). "Xenodens calminechari gen. et sp. nov., a bizarre mosasaurid (Mosasauridae, Squamata) with shark-like cutting teeth from the upper Maastrichtian of Morocco, North Africa". Cretaceous Research. 123 104764. Bibcode:2021CrRes.12304764L. doi:10.1016/j.cretres.2021.104764. S2CID 233567615.
  41. ^ a b Longrich, N. R.; Bardet, N.; Jalil, N.-E.; Pereda-Suberbiola, X.; Schulp, A.; Ghamizi, M. (2025). "New Information on the Morphology and Tooth Replacement of Xenodens calminechari (Squamata: Mosasauridae), a Unique Mosasaurid from the Maastrichtian Phosphates of Morocco". Diversity. 17 (12) 819. doi:10.3390/d17120819.
  42. ^ Sharpe, Henry S.; Powers, Mark J.; Caldwell, Michael W. (2024). "Reassessment of Xenodens calminechari with a discussion of tooth morphology in mosasaurs". The Anatomical Record. 308 (8): 2160–2172. doi:10.1002/ar.25612. ISSN 1932-8486. PMC 12239698.
  43. ^ Houssaye, Alexandra; Rage, Jean-Claude; Bardet, Nathalie; Vincent, Peggy; Amaghzaz, Mbarek; Meslouh, Said (2013). "New highlights about the enigmatic marine snake Palaeophis maghrebianus (Palaeophiidae; Palaeophiinae) from the Ypresian (Lower Eocene) phosphates of Morocco". Palaeontology. 56 (3): 647–661. Bibcode:2013Palgy..56..647H. doi:10.1111/pala.12008.
  44. ^ a b c d Houssaye, Alexandra; Bardet, Nathalie; Rage, Jean-Claude; Suberbiola, Xabier Pereda; Bouya, Baâdi; Amaghzaz, Mbarek; Amalik, Mohamed (2011). "A review of Pachyvaranus crassispondylus Arambourg, 1952, a pachyostotic marine squamate from the latest Cretaceous phosphates of Morocco and Syria". Geological Magazine. 148 (2): 237–249. doi:10.1017/S0016756810000580. ISSN 1469-5081.
  45. ^ a b c d e f g Lomax, Dean R.; Wahl, William R. (2013). "A new specimen of the elasmosaurid plesiosaur Zarafasaura oceanis from the Upper Cretaceous (Maastrichtian) of Morocco" (PDF). Paludicola. 9 (2): 97–109. S2CID 204842778.
  46. ^ a b c d Vincent, Peggy; Bardet, Nathalie; Houssaye, Alexandra; Amaghzaz, Mbarek; Meslouh, Saïd (2013). "New plesiosaur specimens from the Maastrichtian Phosphates of Morocco and their implications for the ecology of the latest Cretaceous marine apex predators". Gondwana Research. 24 (2): 796–805. doi:10.1016/j.gr.2012.11.011. ISSN 1342-937X.
  47. ^ Vincent, Peggy; Bardet, Nathalie; Pereda Suberbiola, Xabier; Bouya, Baâdi; Amaghzaz, Mbarek; Meslouh, Saïd (2011). "Zarafasaura oceanis, a new elasmosaurid (Reptilia: Sauropterygia) from the Maastrichtian Phosphates of Morocco and the palaeobiogeography of latest Cretaceous plesiosaurs". Gondwana Research. 19 (4): 1062–1073. doi:10.1016/j.gr.2010.10.005. ISSN 1342-937X.
  48. ^ Bardet, Nathalie; Jalil, Nour-Eddine; de Lapparent de Broin, France; Germain, Damien; Lambert, Olivier; Amaghzaz, Mbarek (2013). "A giant chelonioid turtle from the Late Cretaceous of Morocco with a suction feeding apparatus unique among tetrapods". PLOS ONE. 8 (7) e63586. Bibcode:2013PLoSO...863586B. doi:10.1371/journal.pone.0063586. PMC 3708935. PMID 23874378.
  49. ^ a b c d Gaffney, E. S.; Tong, H.; Meylan, P. A. (2006). "Evolution of the side-necked turtles: The families Bothremydidae, Euraxemydidae, and Araripemydidae" (PDF). Bulletin of the American Museum of Natural History. 300: 1–318. doi:10.1206/0003-0090(2006)300[1:eotstt]2.0.co;2. hdl:2246/5824. S2CID 85790134.
  50. ^ Tong, H.; Hirayama, R. (2008). "A new species of Argillochelys (Testudines: Cryptodira: Cheloniidae) from the Ouled Abdoun phosphate basin, Morocco". Bulletin de la Société Géologique de France. 179 (6): 623–630. doi:10.2113/gssgfbull.179.6.623.
  51. ^ Nathalie Bardet; Xabier Pereda Suberbiola; Stephane Jouve; Estelle Bourdon; Peggy Vincent; Alexandra Houssaye; Jean-Claude Rage; N.-E Jalil; B. Bouya; M. Amaghzaz (2010). "Reptilian assemblages from the latest Cretaceous – Palaeogene phosphates of Morocco: from Arambourg to present time". Historical Biology. 22 (1–3): 186–199. Bibcode:2010HBio...22..186B. doi:10.1080/08912961003754945. S2CID 128481560.
  52. ^ Lapparent de Broin, France de; Murelaga, Xabier; Vacant, Renaud (October–December 2025). "A chelonioid turtle with a long, upward curved, lower jaw beak, phosphates of Morocco, Paleocene-Eocene transition". Annales de Paléontologie. 111 (4). 102901. doi:10.1016/j.annpal.2025.102901.
  53. ^ Bardet, N.; Jalil, N. E.; De Lapparent De Broin, F.; Germain, D.; Lambert, O.; Amaghzaz, M. (2013). Viriot, Laurent (ed.). "A Giant Chelonioid Turtle from the Late Cretaceous of Morocco with a Suction Feeding Apparatus Unique among Tetrapods". PLOS ONE. 8 (7) e63586. Bibcode:2013PLoSO...863586B. doi:10.1371/journal.pone.0063586. PMC 3708935. PMID 23874378.
  54. ^ a b Gheerbrant, Emmanuel; Sudre, Jean; Iarochene, Mohamed; Moumni, Abdelkader (2001). "First ascertained African "Condylarth" mammals (primitive ungulates: cf. Bulbulodentata and cf. Phenacodonta) from the earliest Ypresian of the Ouled Abdoun Basin, Morocco". Journal of Vertebrate Paleontology. 21 (1): 107–118. doi:10.1671/0272-4634(2001)021[0107:FAACMP]2.0.CO;2. S2CID 85966740.
  55. ^ Gheerbrant, E.; Sudre, J.; Cappetta, H.; Iarochène, M.; Amaghzaz, M.; Bouya, B. (2002). "A new large mammal from the Ypresian of Morocco: Evidence of surprising diversity of early proboscideans" (PDF). Acta Palaeontologica Polonica. 47 (3): 493–506.
  56. ^ Gheerbrant, E. (2009). "Paleocene emergence of elephant relatives and the rapid radiation of African ungulates". Proceedings of the National Academy of Sciences. 106 (26): 10717–10721. Bibcode:2009PNAS..10610717G. doi:10.1073/pnas.0900251106. PMC 2705600. PMID 19549873.
  57. ^ Gheerbrant, Emmanuel (2023-04-27). "Ancestral radiation of paenungulate mammals (Paenungulatomorpha)—new evidence from the Paleocene of Morocco". Journal of Vertebrate Paleontology. 42 (5) e2197971. doi:10.1080/02724634.2023.2197971. ISSN 0272-4634.
  58. ^ Emmanuel Gheerbrant; Arnaud Schmitt; László Kocsis (2018). "Early African fossils elucidate the origin of embrithopod mammals". Current Biology. Online edition. doi:10.1016/j.cub.2018.05.032.
  59. ^ SOLÉ, FLORÉAL; GHEERBRANT, EMMANUEL; AMAGHZAZ, MBAREK; BOUYA, BAÂDI (2009). "Further evidence of the African antiquity of hyaenodontid ('Creodonta', Mammalia) evolution". Zoological Journal of the Linnean Society. 156 (4): 827–846. doi:10.1111/j.1096-3642.2008.00501.x.
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