Ouled Abdoun Basin

Coordinates: 32°53′N 6°55′W / 32.883°N 6.917°W / 32.883; -6.917
Source: Wikipedia, the free encyclopedia.
Ouled Abdoun Basin
Stratigraphic range: Cenomanian-Ypresian
~70.6–25 Ma
The Oulad Abdoun and other major phosphate basins (in yellow) of Morocco
TypeGeological formation
Unit ofSedimentary Basins
Coordinates32°53′N 6°55′W / 32.883°N 6.917°W / 32.883; -6.917
RegionBéni Mellal-Khénifra, Khouribga
Country Morocco
Ouled Abdoun Basin is located in Morocco
Ouled Abdoun Basin
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 (Cenomanian-Turonian) to the Eocene epoch (Ypresian), a period of about 25 million years.[3]


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]


Life restoration of Ouled Abdoun Basin during the late cretaceous: paleofauna roaming on land (left) and sea (right).

Color key
Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
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]

Bony fish

Bony fish from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Acipenseridae[5] 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[6] E. cattoi Maastrichtian A pycnodontiform.
Mawsoniidae[7] Indeterminate Maastrichtian A giant mawsoniid coelacanth measuring 3.65–5.52 metres (12.0–18.1 ft) long.
Phacodus[8] P. punctatus var. africanus Maastrichtian A pycnodontiform.
Plethodidae[9] Indeterminate Maastrichtian Youngest occurence of plethodid fish.
Pycnodontidae Indeterminate Maastrichtian
Pseudoegertonia P. sp. Maastrichtian
Serrasalmimus[6] S. secans Thanetian A carnivorous pycnodontiform.
Stephanodus S. libycus Maastrichtian
Stratodus S. apicalis Maastrichtian An alepisauroform.


Sharks from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Abdounia[10] A. africana Selandian A requiem shark.
Casieria C. maghrebiana Selandian
Chiloscyllium C. salvani. Selandian A carpet shark.
Cretalamna C. appendiculata Maastrichtian
C. maroccana Maastrichtian
Delpitoscyllium D. africanum Selandian
Eostegostoma E. sp. Selandian A carpet shark.
Foumtizia F. abdouni Selandian
Galeorhinus G. mesetaensis Selandian
Ginglymostoma G. chenanei Selandian
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
Scyliorhinus S. ptychtus Selandian
S. entomodon Selandian
S. sulcidens Selandian
Serratolamna S. serrata Maastrichtian
Serratolamna teeth
S. khderii Maastrichtian
Squalicorax S. pristodontus Maastrichtian
S. bassanii Maastrichtian
S. africanus Maastrichtian
S. microserratus Maastrichtian
S. benguerirensis Maastrichtian
Triakis T. antunesi Selandian

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 stringray.
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.
Rhombodus spines
R. microdus Maastrichtian
R. meridionalis Maastrichtian
Schizorhiza S. stromeri Maastrichtian A sclerorhynchoid skate which resembled a sawfish.


Crocodylomorphs from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Arambourgisuchus[11] A. khouribgaensis Thanetian A dyrosaurid.
Atlantosuchus A. coupatezi Danian A dyrosaurid.
Chenanisuchus C. lateroculi Thanetian The most basal known dyrosaurid.
Dyrosaurus P. maghribensis Ypresian A dyrosaur.
Maroccosuchus[12] M. zennaroi Ypresian A tomistomine crocodylian.
Ocepesuchus[13] O. eoafricanus Maastrichtian A gavialoid crocodylian.


Lepidosaurs from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Mosasaurus M. beaugei Maastrichtian A large mosasaurine mosasaur.
Eremiasaurus E. heterodontus Maastrichtian A mosasaurine mosasaur with unusual cutting dentition.
Halisaurus H. arambourgi Maastrichtian A halisaurine mosasaur.
Globidens G. phosphaticus Maastrichtian A mosasaurine mosasaur with specialized crushing teeth.
G. simplex Maastrichtian
G. aegyptiacus Maastrichtian Now referred to its own genus, Igdamanosaurus.
Igdamanosaurus I. aegyptiacus Maastrichtian
Carinodens C. minalmamar Maastrichtian A small mosasaurine mosasaur with crushing teeth.
Platecarpus P. ptychodon Maastrichtian A nomen dubium. Now referred to its own genus, Gavialimimus.
Gavialimimus[14] G. almaghribensis Maastrichtian An unusual longirostrine plioplatecarpine mosasaur.
Prognathodon P. currii Maastrichtian A large mosasaurine mosasaur.
P. giganteus Maastrichtian
Thalassotitan[15] T. atrox Maastrichtian A large-bodied mosasaurine with powerful jaws, similar to P. currii.
Stelladens[16] S. mysteriosus Maastrichtian A medium-sized mosasaurine with distinct, unique teeth structure.
Pluridens[17] P. serpentis Maastrichtian A large halisaurine mosasaur, also known from Niger and Nigeria.
Xenodens[18] X. calminechari Maastrichtian A small mosasaurine mosasaur with unusual, shark-like teeth.
Pachyvaranus P. crassispondylus Maastrichtian A pachyvaranid
Palaeophis[19] P. maghrebianus Maastrichtian?–Ypresian A palaeophiid marine snake.


Plesiosaurs from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Zarafasaura Z. oceanis Maastrichtian An elasmosaurid plesiosaur with an unusually short skull.


Turtles from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Alienochelys[20] A. selloumi Maastrichtian A sea turtle with crushing jaws.
Araiochelys[21] A. hirayamai
Argillochelys[22] A. africana Ypresian A pancheloniid sea turtle.
Bothremys[21] B. kellyi Danian-Thanetian
B. maghrebiana
Brachyopsemys B. tingitana Danian A sandownid sea turtle
Euclastes E. wielandi Danian
Labrostochelys[21] L. galkini Danian
Ocepechelon[23] O. bouyai Maastrichtian A giant protostegid sea turtle with an unusual pipette-like snout.
Puppigerus P. camperi Ypresian
Rhothonemys[21] R. brinkmani Danian
Tasbacka T. ouledabdounensis Danian
Taphrosphys T. ippolitoi Danian
Ummulisani U. rutgersensis Ypresian


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


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]

Dinosaurs from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Abelisauridae[25] Indeterminate Sidi Daoui Maastrichtian right metatarsal II A mature specimen but very small for an abelisaurid, about 2.6 m (8.5 ft) long
Abelisauridae[25] 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.
Ajnabia[26] 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.
Chenanisaurus[27] C. barbaricus
  • Sidi Chennane
  • Sidi Daoui
  • anterior end of left dentary
  • 2 premaxillary teeth
  • a maxillary tooth
A large abelisaurid theropod.
Titanosauria Indeterminate Maastrichtian A new taxon, currently unnamed.
Dasornis D. tolapica Selandian A pelagornithid, or pseudotooth bird.
D. emuinus Ypresian
Lithoptila[28] L. abdounensis Thanetian A seabird related to modern tropicbirds.


Mammals from the Khouribga Phosphates
Genus Species Location Time Material Notes Images
Abdounodus[29] A. hamdii Thanetian-Ypresian
Daouitherium[30] D. rebouli Ypresian An early proboscidean.
Eritherium[31] E. azzouzorum Thanetian An early proboscidean.
Hadrogeneios[32] H. phosphaticus Selandian-Thanetian
Ocepeia[29] O. daouiensis Selandian

O. grandis Thanetian
Phosphatherium P. escuilliei Selandian An early proboscidean.
Boualitomus B. marocanensis Selandian A hyaenodont.
Lahimia[33] L. selloumi Selandian A hyaenodont.
Stylolophus[34] S. minor Ypresian An embrithopod.
S. major Ypresian

See also


  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. ^ Martill, D. M. (2023). "A sturgeon (Actinopterygii, Acipenseriformes) from the Upper Cretaceous of Africa". Cretaceous Research. 148. 105546. Bibcode:2023CrRes.14805546M. doi:10.1016/j.cretres.2023.105546. S2CID 257863907.
  6. ^ 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. doi:10.1038/s41598-017-06792-x. ISSN 2045-2322. PMC 5533729.
  7. ^ 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. doi:10.1016/j.cretres.2021.104768. S2CID 233551515.
  8. ^ 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. doi:10.1016/j.cretres.2021.105111. ISSN 0195-6671.
  9. ^ 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: 105673. doi:10.1016/j.cretres.2023.105673. ISSN 0195-6671.
  10. ^ 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.
  11. ^ 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.
  12. ^ 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.
  13. ^ 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.
  14. ^ 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. doi:10.1080/14772019.2020.1818322. ISSN 1477-2019.
  15. ^ 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. doi:10.1016/j.cretres.2022.105315. S2CID 251821884.
  16. ^ 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. ISSN 2813-6284.
  17. ^ 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" (PDF). Cretaceous Research. 126: Article 104882. doi:10.1016/j.cretres.2021.104882.
  18. ^ 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" (PDF). Cretaceous Research. 123: Article 104764. doi:10.1016/j.cretres.2021.104764. S2CID 233567615.
  19. ^ 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. doi:10.1111/pala.12008.
  20. ^ 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.
  21. ^ 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.
  22. ^ 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.
  23. ^ 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.
  24. ^ 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.
  25. ^ 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: 105677. doi:10.1016/j.cretres.2023.105677.
  26. ^ 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.
  27. ^ 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. doi:10.1016/j.cretres.2017.03.021.
  28. ^ 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.
  29. ^ 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.
  30. ^ 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.
  31. ^ 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.
  32. ^ Gheerbrant, Emmanuel (2023-04-27). "Ancestral radiation of paenungulate mammals (Paenungulatomorpha)—new evidence from the Paleocene of Morocco". Journal of Vertebrate Paleontology. doi:10.1080/02724634.2023.2197971. ISSN 0272-4634.
  33. ^ 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.
  34. ^ 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.

External links