Kem Kem Group

The Kem Kem Group (commonly known as the Kem Kem beds^[2]) is a geological group in the Kem Kem region of eastern Morocco, whose strata date back to the Cenomanian stage of the Late Cretaceous. Its strata are subdivided into two geological formations, with the lower Ifezouane Formation and the upper Aoufous Formation used for the strata on the eastern side of the Atlas Mountains (Tinghir), with the Gara Sbaa Formation and Douira Formation used in the southern Tafilalt region.^[3] It is exposed on an escarpment along the Algeria–Morocco border.

Kem Kem Group
Stratigraphic range: Cenomanian[1] ~98–94 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Type	Geological group
Sub-units	Douira Formation, Gara Sbaa Formation
Underlies	Cenomanian-Turonian limestone platform (Akrabou Formation)
Lithology
Primary	Sandstone
Location
Coordinates	32°50′N 4°50′W
Approximate paleocoordinates	18.8°N 4.1°W / 18.8; -4.1
Region	Er Rachidia, Tafilalt
Country	Morocco
Extent	central and eastern Morocco north and south of the Pre-African Trough
Kem Kem Group (Morocco)

The unit unconformably overlies Paleozoic marine units of Cambrian, Silurian and Devonian ages and is itself capped by limestone platform rock of Cenomanian-Turonian age. It primarily consists of freshwater and estuarine deltaic deposits. The lower Gara Sbaa Formation primarily consists of fine and medium grained sandstone, while the Douira Formation consists of fining-upward, coarse-to-fine grained sandstones intercalated with siltstones, variegated mudstones, and occasional thin gypsiferous evaporites.^[2]

Dinosaur remains are among the fossils that have been recovered from the group.^[1] Recent fossil evidence in the form of isolated large abelisaurid bones and comparisons with other similarly aged deposits elsewhere in Africa indicates that the fauna of the Kem Kem Group (specifically in regard to the numerous predatory theropod dinosaurs) may have been mixed together due to the harsh and changing geology of the region, when in reality they would likely have preferred separate habitats and likely would have been separated by millions of years.^[4]

Although preserving a freshwater habitat located near a river delta (with some estuarine influence that increased over time as the sea level rose), the Kem Kem deposits were quickly submerged by the sea during the Cenomanian-Turonian boundary event, and are thus overlaid by the marine deposits of the younger latest Cenomanian and early-mid Turonian-aged Akrabou Formation, which was formerly also considered a member of the Kem Kem Group, but has been differentiated from it in more recent studies due to their differing paleoenvironments.^[2][5]

Vertebrate paleofauna

Cartilaginous fish

Cartilaginous fish
Genus	Species	Location	Material	Notes	Images
Acrodontidae indet.[2]	Indeterminate			Members of Hybodontoidea
Bahariyodon[2]	B. bartheli			A member of Hybodontoidea
Cenocarcharias[2]	C. tenuiplicatus		One tooth[2]	A member of the family Cretoxyrhinidae
Distobatus[2]	D. nutiae			A member of Hybodontoidea
Haimirichia[2]	H. amonensis		One tooth[2]	A mackerel shark
Marckgrafia[2]	M. lybica		13 teeth[2]	A member of Batoidea
Onchopristis	O. numida[6]			A rajiform ray[7]	Rostrum and teeth fossils from Onchopristis
Peyeria[2]	P. libyca		Three teeth[2]	A sawfish. Might be a junior synonym of Onchopristis numida.
Tribodus[2]	T. sp.			A member of Hybodontoidea

Ray-finned fish

Ray-finned fish
Genus	Species	Location	Material	Notes	Images
Adrianaichthys[2]	A. pankowskii		Isolated scales[8] and two skulls[9]	A member of Lepisosteiformes. Originally described as a species of Lepidotes, but subsequently transferred to a separate genus.[10]	Life restoration of Aidachar
Afrocascudo[11]	A. saharaensis			A neopterygiian fish, either an ancient loricariid catfish or a juvenile obaichthyid lepisosteiform.[12]
Agassizilia[13]	A. erfoudina			Possibly a member of the family Pycnodontidae.
Agoultichthys[2]	A. chattertoni			A long-bodied member of Actinopterygii of uncertain phylogenetic placement. Might be a member of the family Macrosemiidae[14] or Ophiopsiellidae.[15]
Aidachar	A. pankowskii			A member of Ichthyodectiformes
Bartschichthys[2]	B. sp.		Isolated pinnulae (spines that support each dorsal finlet)[2]	A cladistian
Bawitius	cf. B. sp.		Isolated scales and jaw fragments[8]	A cladistian
Calamopleurus[2]	C. africanus		A partial skull[2]	A member of Amiiformes
Concavotectum[2]	C. moroccensis			A member of Tselfatiiformes
Dentilepisosteus[2]	D. kemkemensis			A member of Lepisosteiformes
Diplomystus[2]	D. sp.			A deep-bodied teleost belonging to the group Clupeomorpha
Diplospondichthys[2]	D. moreaui			A member of Actinopterygii of uncertain phylogenetic placement, possibly a teleost
Erfoudichthys[2]	E. rosae		Isolated skull[2]	A small-bodied teleost of unknown affinity
Neoproscinetes[13]	N. africanus			A member of the family Pycnodontidae
Obaichthys	O. africanus		Isolated scales[8]	A member of Lepisosteiformes
Oniichthys	O. falipoui		Near complete skeleton including skull[8]	A member of Lepisosteiformes
Palaeonotopterus[2]	P. greenwoodi			A member of Osteoglossomorpha
Serenoichthys[2]	S. kemkemensis		Several articulated skeletons[2]	A small cladistian
Spinocaudichthys[2]	S. oumtkoutensis			An elongate freshwater acanthomorph
Stromerichthys	S. aethiopicus
Sudania[2]	S. sp.		An isolated pinnula[2]	A cladistian

Lobe-finned fish

Lobe-finned fish
Genus	Species	Location	Material	Notes	Images
Arganodus	A. tiguidiensis			A lungfish	Life restoration of Axelrodichthys The Queensland Lungfish, the only living member of Neoceratodus
Axelrodichthys[16]	A.? lavocati			A mawsoniid coelacanth; this species was previously assigned to Mawsonia, and its generic assignment is still not certain[17]
Neoceratodus	N. africanus			A lungfish

Amphibians

Amphibians
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Anura indet.[18]	Indeterminate		Douira Formation	Incomplete left ilium
Cretadhefdaa[18]	C. taouzensis		Douira Formation	Posterior portion of the skull, incomplete squamosal, incomplete maxilla, three incomplete presacral vertebrae, one incomplete sacral vertebra	A neobatrachian frog with possible hyloid affinities.
cf. Kababisha[19]	Indeterminate				A salamander belonging to the family Sirenidae
?Neobatrachia indet.[18]	Indeterminate		Douira Formation	Incomplete humerus	A frog, possibly a member of Ranoidea.
Oumtkoutia[19]	O. anae				A frog belonging to the family Pipidae

Lizards and snakes

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.

Lizards and snakes reported from the Continental Red Beds
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Bicuspidon	B. hogreli[20]				A polyglyphanodontid lizard.
Jeddaherdan[21]	J. aleadonta			Partial mandible with teeth.	An iguanian belonging to the group Acrodonta, possibly a relative of the uromastycine agamids. Argued by Vullo et al. (2022) to actually come from Quaternary beds, and to be based on a fossil material of a member of the genus Uromastyx.[22]
Lapparentophis	L. ragei[23]			Two isolated trunk vertebrae	An early snake.
Madtsoiidae indet.[19]	Indeterminate			Vertebrae[2]	An early snake.
?Nigerophiidae indet.[19]	Indeterminate			Dorsal vertebrae[2]	An early snake.
Norisophis[24]	N. begaa[24]			One posterior and two mid-trunk vertebrae	A stem-snake.
	Indeterminate[24]			A mid-trunk vertebra
Simoliophis[19]	cf. S. libycus			Vertebrae[2]	An early snake.

Plesiosaurs

Plesiosaurs
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Leptocleididae cf. Leptocleidus[25]	indeterminate

Turtles

Turtles reported from the Continental Red Beds
Genus	Species	Location	Stratigraphic position	Material	Notes
Dirqadim	D. schaefferi				A Euraxemydid
Galianemys	G. emringeri				A Cearachelyin
	G. whitei
Hamadachelys	H. escuilliei

Crocodylomorphs

A tooth enamel identified as cf. Sarcosuchus was discovered from the Ifezouane Formation.^[26]

Crocodylomorphs reported from the Continental Red Beds
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Aegisuchus	A. witmeri			"Partial braincase of a large individual with skull roof, temporal, and occipital regions."[27]	An aegyptosuchid that may be a synonym of Laganosuchus.[2]	Aegisuchus Araripesuchus Elosuchus cherifiensis Laganosuchus
Antaeusuchus[28]	A. taouzensis			Paired mandibles and a partial right mandible	A peirosaurid.
Araripesuchus	A. rattoides		Douira Formation
Elosuchus	E. cherifiensis		Gara Sbaa Formation Douira Formation		An elosuchid. The material may represent two different species.[2]
Hamadasuchus	H. rebouli		Gara Sbaa Formation Douira Formation?		A peirosaurid.
Kemkemia	K. auditorei	Errachidia Province, Morocco[29]		Known from an isolated caudal vertebra.[29]	Initially thought to be a neotheropod,[29] but subsequently discovered to be an indeterminate crocodyliform.[30]
Laganosuchus	L. thaumastos				A stomatosuchid.
Lavocatchampsa[31]	Lavocatchampsa sigogneaurusselae			Anterior portion of a rostrum with mandible, with an almost complete dentition[31]	A candidodontid notosuchian.[31]

Dinosaurs

Indeterminate lithostrotian remains once misattributed to the Titanosauridae are present in the province of Ksar-es-Souk, Morocco.^[1]

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.

Ornithischians

Ornithischians reported from the Continental Red Beds
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Ankylosauria[2][32]	Indeterminate		Douira Formation	An isolated tooth.[2]	A probable ankylosaur[32]
Ornithopoda[33]	Indeterminate		Douira Formation	A large, clover-shaped, three-toed footprint.[2]	Comparable in size and shape to tracks typically attributed to Iguanodon.[33]

Sauropods

Sauropods reported from the Continental Red Beds
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Rebbachisaurus	R. garasbae	Ksar-es-Souk province, Morocco.[1]	Gara Sbaa Formation		A rebbachisaurid.
Somphospondyli[34]	Indeterminate			Anterior dorsal vertebra, partial right ischium[34]	The vertebra might belong to a basal titanosaurian, possibly distinct from Aegyptosaurus and Paralititan.[34] The ischium is not identifiable beyond Somphospondyli; it preserves numerous grooves and pits which might be feeding traces left by a very large non-avian theropod.[34]
Titanosauria[35][2]	Indeterminate		Ifezouane Formation Douira Formation	Isolated teeth, caudal vertebrae, a partial humerus, a tarsal bone and the proximal end of an ulna.[2]	Fossil material of one or more titanosaurian sauropods. Some fossils are indicative of large body size comparable to Paralititan stromeri.[2]

Theropods

Theropods reported from the Continental Red Beds
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Abelisauridae[36]	Indeterminate			Isolated teeth.[36][37] Partial right femur.[4] Partial right maxilla.[2] A cervical vertebra.[38] Left maxilla.[2]	Abelisaurid material belonging to one or two distinct taxa.[2]	Carcharodontosaurus saharicus Deltadromeus agilis Rebbachisaurus garasbae Rugops primus Spinosaurus aegyptiacus reconstruction
Carcharodontosaurus[1]	C. saharicus[1]	Ksar-es-Souk province, Morocco.[1]	Douira Formation	Partial skull, including braincase, nasals, postorbitals, jugals, left lacrimal and right maxilla with most teeth.[39]	A carcharodontosaurid theropod.
Carcharodontosauridae[40]	Indeterminate	Southeast of Taouz, Errachidia Province	Ifezouane Formation	partial maxilla and partial jugal	A carcharodontosaurid theropod different from C. saharicus
Deltadromeus	D. agilis		Gara Sbaa Formation	"Partial skeleton, isolated limb elements."[41]	A noasaurid ceratosaurian or possible neovenatorid carnosaur. May be synonymous with Bahariasaurus.
Dromaeosauridae[36]	Indeterminate			Isolated teeth.[36]	Originally described as teeth of indeterminate dromaeosaurids. Hendrickx et al. (2024) reinterpreted this fossil material as teeth of abelisauroid theropods, including noasaurids and juvenile abelisaurids.[37]
cf. Elaphrosaurus	Indeterminate	Ksar-es-Souk province, Morocco.[1]			Fossils previously referred to cf. Elaphrosaurus are actually indeterminate theropod remains.
Noasauridae[38]	Indeterminate			An anterior cervical vertebra[38] Isolated teeth[37]
"Osteoporosia"[42]	"O. gigantea"[42]			A tooth and a possible neural arch from another specimen.[42]	A theropod, possibly synonymous with Sauroniops.[43]
Saurischia[44]	Indeterminate			An isolated cervical vertebra.[44]	An indeterminate saurischian.
Sauroniops[45]	S. pachytholus		Ifezouane Formation	"An isolated and almost complete left frontal,[46] and a possible tooth and neural arch from two other specimens."[43]	A carcharodontosaurid distinct from Carcharodontosaurus.[45][46]
Spinosaurus	S. aegyptiacus	Ksar-es-Souk province, Morocco.[1]	Douira Formation	Partial skeleton, including parts of the skull, neck, torso, and most of the tail and hind limbs. [47] Numerous isolated bones.

Pterosaurs

Pterosaurs of the Kem Kem Beds
Genus	Species	Location	Stratigraphic position	Abundance	Notes	Images
Afrotapejara[48]	A. zouhri[48]	Takmout	Ifezouane Formation	A fragment of bone interpreted as a fragment of anterior mandibular symphysis,[49] and additional jaw fragments that pertain to the rostrum as well as indeterminate jaw fragments.[3]	A tapejarid pterosaur. Originally believed to belong to either the family Thalassodromidae[50] or an additional specimen of Alanqa saharica.[51]	Afrotapejara zouhri Akharhynchus martilli Alanqa saharica Anhanguera Coloborhynchus Leptostomia Ornithocheirus
Akharhynchus[52]	A. martilli	Tafilalt	Ifezouane Formation	A fragment of the anterior part of the premaxillae	A tropeognathine anhanguerian.
Alanqa[50]	A. saharica[50]		Ifezouane Formation	The holotype is a mandibular symphysis, of different parts of the jaw	A pterosaur of uncertain phylogenetic placement, probably an azhdarchid.[3]
Anhanguera[53]	A. cf. piscator[53]		upper Ifezouane Formation	Partial mandibular symphysis[53]
Apatorhamphus[54]	A. gyrostega[54]		Ifezouane Formation	Partial rostrum and mandible, with additional referred jaw fragments[3]	A possible chaoyangopterid azhdarchoid pterosaur.[54] Originally believed to be a possible pteranodontid,[50] a possible dsungaripterid,[55] a possible non-azhdarchid azhdarchoid or nyctosaurid,[55] or a specimen of Alanqa saharica.[51]
Azhdarchidae indet.[55]	Indeterminate[55]			Three middle cervical vertebrae.[55][50]	Averianov (2014) considered these vertebrae to pertain to Alanqa saharica,[51] although the vertebrae may be indicative of two taxa.[55]
Coloborhynchus[53]	C. sp. A.[53]	Hassi El Begaa		Premaxillae fragment[53]	Possibly a specimen of Nicorhynchus fluviferox.[56]
Leptostomia[57]	L. begaaensis[57]	Aferdou N’ Chaft	upper Ifezouane Formation	Partial rostrum and partial mandibular synthesis[57]	A small, long-beaked pterosaur, likely a member of Azhdarchoidea.[57]
Nicorhynchus[56]	N. fluviferox[58][56]	Possibly Aferdou N’Chaft, Hassi El Begaa[56]	Ifezouane Formation	An anterior portion of the rostrum.[56]	Originally described as a species of Coloborhynchus[58] but subsequently transferred to the genus Nicorhynchus.
Ornithocheirus[53]	O. cf. simus.[53]		upper Ifezouane Formation	Premaxillae fragment[53]
Siroccopteryx[59]	S. moroccensis[59]			Anterior part of a rostrum	Classified by some authors as a species belonging to the genus Coloborhynchus.[50]
Xericeps	X. curvirostra	Aferdou N'Chaft	Ifezouane Formation	Mandibular symphysis and partial mandible[3]	An indeterminate azhdarchoid, possibly a chaoyangopterid.[3]

See also

• Earth sciences portal
• Paleontology portal
• Dinosaurs portal

• Aoufous Formation, which lies within the Kem Kem Beds
• List of dinosaur-bearing rock formations

References

1. Weishampel, David B; et al. (2004). "Dinosaur distribution (Late Cretaceous, Africa)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria, 2nd, Berkeley: University of California Press. Pp. 604-605. ISBN 0-520-24209-2.
2. Ibrahim, N.; Sereno, P.C.; Varricchio, D.J.; Martill, D.M.; Dutheil, D.B.; Unwin, D.M.; Baidder, L.; Larsson, H.C.E.; Zouhri, S.; Kaoukaya, A. (2020). "Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco". ZooKeys (928): 1–216. Bibcode:2020ZooK..928....1I. doi:10.3897/zookeys.928.47517. PMC 7188693. PMID 32362741.
3. Smith, Roy E.; Ibrahim, Nizar; Longrich, Nicholas; Unwin, David M.; Jacobs, Megan L.; Williams, Cariad J.; Zouhri, Samir; Martill, David M. (2023-02-04). "The pterosaurs of the Cretaceous Kem Kem Group of Morocco". PalZ. 97 (3): 519–568. Bibcode:2023PalZ...97..519S. doi:10.1007/s12542-022-00642-6. ISSN 1867-6812. S2CID 256608633.
4. Alfio Alessandro Chiarenza; Andrea Cau (2016). "A large abelisaurid (Dinosauria, Theropoda) from Morocco and comments on the Cenomanian theropods from North Africa". PeerJ. 4: e1754. doi:10.7717/peerj.1754. PMC 4782726. PMID 26966675.
5. Cavin, L.; Tong, H.; Boudad, L.; Meister, C.; Piuz, A.; Tabouelle, J.; Aarab, M.; Amiot, R.; Buffetaut, E.; Dyke, G.; Hua, S.; Le Loeuff, J. (2010-07-01). "Vertebrate assemblages from the early Late Cretaceous of southeastern Morocco: An overview". Journal of African Earth Sciences. 57 (5): 391–412. doi:10.1016/j.jafrearsci.2009.12.007. ISSN 1464-343X.
6. Greenfield, T. (2021). "Corrections to the nomenclature of sawskates (Rajiformes, Sclerorhynchoidei)". Bionomina. 22 (1): 39–41. doi:10.11646/bionomina.22.1.3. S2CID 239067365.
7. Villalobos-Segura, Eduardo; Kriwet, Jürgen; Vullo, Romain; Stumpf, Sebastian; Ward, David J; Underwood, Charlie J (2021-10-01). "The skeletal remains of the euryhaline sclerorhynchoid †Onchopristis (Elasmobranchii) from the 'Mid'-Cretaceous and their palaeontological implications". Zoological Journal of the Linnean Society. 193 (2): 746–771. doi:10.1093/zoolinnean/zlaa166. ISSN 0024-4082.
8. Cavin, Lionel; Boudad, Larbi; Tong, Haiyan; Läng, Emilie; Tabouelle, Jérôme; Vullo, Romain (2015). "Taxonomic Composition and Trophic Structure of the Continental Bony Fish Assemblage from the Early Late Cretaceous of Southeastern Morocco". PLOS ONE. 10 (5): e0125786. Bibcode:2015PLoSO..1025786C. doi:10.1371/journal.pone.0125786. ISSN 1932-6203. PMC 4446216. PMID 26018561.
9. Forey, Peter L.; López-Arbarello, Adriana; MacLeod, Norman (2011). "A New Species of Lepidotes (Actinopterygii: Semiontiformes) from the Cenomanian (Upper Cretaceous) of Morocco" (PDF). Palaeontologia Electronica. 14 (1): 7A–12.
10. François J. Meunier; René-Paul Eustache; Didier Dutheil; Lionel Cavin (2016). "Histology of ganoid scales from the early Late Cretaceous of the Kem Kem beds, SE Morocco: systematic and evolutionary implications". Cybium. 40 (2): 121–132. doi:10.26028/cybium/2016-402-003.
11. Brito, P. M.; Dutheil, D. B.; Gueriau, P.; Keith, P.; Carnevale, G.; Britto, M.; Meunier, F. J.; Khalloufi, B.; King, A.; de Amorim, P. F.; Costa, W. J. E. M. (2024). "A saharan fossil and the dawn of Neotropical armoured catfishes in Gondwana". Gondwana Research. 132: 103–112. Bibcode:2024GondR.132..103B. doi:10.1016/j.gr.2024.04.008.
12. Britz, R.; Pinion, Amanda K.; Kubicek, Kole M.; Conway, Kevin W. (2024). "Comment on "A Saharan fossil and the dawn of Neotropical armoured catfishes in Gondwana" by Brito et al". Gondwana Research. 133: 267–269. Bibcode:2024GondR.133..267B. doi:10.1016/j.gr.2024.06.014.
13. Samuel L.A. Cooper; David M. Martill (2020). "A diverse assemblage of pycnodont fishes (Actinopterygii, Pycnodontomorpha) from the mid-Cretaceous, continental Kem Kem beds of South-East Morocco". Cretaceous Research. 112: Article 104456. doi:10.1016/j.cretres.2020.104456. S2CID 216214083.
14. Alison M. Murray; Mark V. H. Wilson (2009). "A new Late Cretaceous macrosemiid fish (Neopterygii, Halecostomi) from Morocco, with temporal and geographical range extensions for the family". Palaeontology. 52 (2): 429–440. Bibcode:2009Palgy..52..429M. doi:10.1111/j.1475-4983.2009.00851.x. S2CID 128548868.
15. Martin Ebert (2018). "Cerinichthys koelblae, gen. et sp. nov., from the Upper Jurassic of Cerin, France, and its phylogenetic setting, leading to a reassessment of the phylogenetic relationships of Halecomorphi (Actinopterygii)". Journal of Vertebrate Paleontology. 38 (1): e1420071. Bibcode:2018JVPal..38E0071E. doi:10.1080/02724634.2017.1420071. S2CID 89886438.
16. Fragoso, L.G.C.; Brito, P.; Yabumoto, Y. (2019). "Axelrodichthys araripensis Maisey, 1986 revisited". Historical Biology. 31 (10): 1350–1372. doi:10.1080/08912963.2018.1454443. S2CID 89795160.
17. Cavin, L.; Cupello, C.; Yabumoto, Y.; Fragoso, L.G.C.; Deesri, U.; Brito, P.M. (2019). "Phylogeny and evolutionary history of mawsoniid coelacanths" (PDF). Bulletin of the Kitakyushu Museum of Natural History and Human History, Series A. 17: 3–13.
18. Lemierre, A.; Blackburn, D. C. (2022). "A new genus and species of frog from the Kem Kem (Morocco), the second neobatrachian from Cretaceous Africa". PeerJ. 10: e13669. doi:10.7717/peerj.13699. PMC 9291016. PMID 35860040.
19. Jean-Claude Rage; Didier B. Dutheil (2008). "Amphibians and squamates from the Cretaceous (Cenomanian) of Morocco - A preliminary study, with description of a new genus of pipid frog". Palaeontographica Abteilung A. 285 (1–3): 1–22. Bibcode:2008PalAA.285....1R. doi:10.1127/pala/285/2008/1.
20. Romain Vullo; Jean-Claude Rage (2018). "The first Gondwanan borioteiioid lizard and the mid-Cretaceous dispersal event between North America and Africa". The Science of Nature. 105 (11–12): Article 61. Bibcode:2018SciNa.105...61V. doi:10.1007/s00114-018-1588-3. PMID 30291449. S2CID 52924052.
21. Sebastián Apesteguía; Juan D. Daza; Tiago R. Simões; Jean Claude Rage (2016). "The first iguanian lizard from the Mesozoic of Africa". Royal Society Open Science. 3 (9): 160462. Bibcode:2016RSOS....360462A. doi:10.1098/rsos.160462. PMC 5043327. PMID 27703708.
22. Romain Vullo; Salvador Bailon; Yannicke Dauphin; Hervé Monchot; Ronan Allain (2022). "A reappraisal of Jeddaherdan aleadonta (Squamata: Acrodonta), the purported oldest iguanian lizard from Africa". Cretaceous Research. 143. 105412. doi:10.1016/j.cretres.2022.105412. S2CID 253349389.
23. Romain Vullo (2019). "A new species of Lapparentophis from the mid-Cretaceous Kem Kem beds, Morocco, with remarks on the distribution of lapparentophiid snakes". Comptes Rendus Palevol. 18 (7): 765–770. Bibcode:2019CRPal..18..765V. doi:10.1016/j.crpv.2019.08.004. S2CID 210297438.
24. Catherine G. Klein; Nicholas R. Longrich; Nizar Ibrahim; Samir Zouhri; David M. Martill (2017). "A new basal snake from the mid-Cretaceous of Morocco" (PDF). Cretaceous Research. 72: 134–141. Bibcode:2017CrRes..72..134K. doi:10.1016/j.cretres.2016.12.001.
25. Bunker, G.; Martill, D. M.; Smith, R.; Zourhi, S.; Longrich, N. (2022). "Plesiosaurs from the fluvial Kem Kem Group (mid-Cretaceous) of eastern Morocco and a review of non-marine plesiosaurs". Cretaceous Research. 140: Article 105310. Bibcode:2022CrRes.14005310B. doi:10.1016/j.cretres.2022.105310. S2CID 250964381.
26. Amiot, Romain; Wang, Xu; Lécuyer, Christophe; Buffetaut, Eric; Boudad, Larbi; Cavin, Lionel; Ding, Zhongli; Fluteau, Frédéric; Kellner, Alexander W.A.; Tong, Haiyan; Zhang, Fusong (2010). "Oxygen and carbon isotope compositions of middle Cretaceous vertebrates from North Africa and Brazil: Ecological and environmental significance". Palaeogeography, Palaeoclimatology, Palaeoecology. 297 (2): 439–451. Bibcode:2010PPP...297..439A. doi:10.1016/j.palaeo.2010.08.027.
27. Casey M. Holliday; Nicholas M. Gardner (2012). "A New Eusuchian Crocodyliform with Novel Cranial Integument and Its Significance for the Origin and Evolution of Crocodylia". PLOS ONE. 7 (1): e30471. Bibcode:2012PLoSO...730471H. doi:10.1371/journal.pone.0030471. PMC 3269432. PMID 22303441.
28. Cecily S. C. Nicholl; Eloise S. E. Hunt; Driss Ouarhache; Philip D. Mannion (2021). "A second peirosaurid crocodyliform from the Mid-Cretaceous Kem Kem Group of Morocco and the diversity of Gondwanan notosuchians outside South America". Royal Society Open Science. 8 (10): Article ID 211254. Bibcode:2021RSOS....811254N. doi:10.1098/rsos.211254. PMC 8511751. PMID 34659786.
29. Cau, Andrea; Maganuco, Simone (2009). "A new theropod dinosaur, represented by a single unusual caudal vertebra from the Kem Kem Beds (Cretaceous) of Morocco". Atti Soc. it. Sci. nat. Museo civ. Stor. nat. Milano 150 (II): 239–257.
30. Lio, G., Agnolin, F., Cau, A. and Maganuco, S. (2012). "Crocodyliform affinities for Kemkemia auditorei Cau and Maganuco, 2009, from the Late Cretaceous of Morocco." Atti della Società Italiana di Scienze Naturali e del Museo di Storia Naturale di Milano, 153 (I), s. 119–126.
31. Jeremy E. Martin; France De Lapparent De Broin (2016). "A miniature notosuchian with multicuspid teeth from the Cretaceous of Morocco" (PDF). Journal of Vertebrate Paleontology. 36 (6): e1211534. Bibcode:2016JVPal..36E1534M. doi:10.1080/02724634.2016.1211534. S2CID 89354184.
32. Rozadilla, Sebastián; Agnolín, Federico; Manabe, Makoto; Tsuihiji, Takanobu; Novas, Fernando E. (September 2021). "Ornithischian remains from the Chorrillo Formation (Upper Cretaceous), southern Patagonia, Argentina, and their implications on ornithischian paleobiogeography in the Southern Hemisphere". Cretaceous Research. 125: 104881. Bibcode:2021CrRes.12504881R. doi:10.1016/j.cretres.2021.104881. ISSN 0195-6671.
33. Nizar Ibrahim; David J. Varricchio; Paul C. Sereno; Jeff A. Wilson; Didier B. Dutheil; David M. Martill; Lahssen Baidder; Samir Zouhri (2014). "Dinosaur footprints and other ichnofauna from the Cretaceous Kem Kem beds of Morocco". PLOS ONE. 9 (6): e90751. Bibcode:2014PLoSO...990751I. doi:10.1371/journal.pone.0090751. PMC 3946209. PMID 24603467.
34. Matthew C. Lamanna; Yoshikazu Hasegawa (2014). "New titanosauriform sauropod dinosaur material from the Cenomanian of Morocco: implications for paleoecology and sauropod diversity in the Late Cretaceous of North Africa" (PDF). Bulletin of Gunma Museum of Natural History. 18: 1–19.
35. Nizar Ibrahim; Cristiano Dal Sasso; Simone Maganuco; Matteo Fabbri; David M. Martill; Eric Gorscak; Matthew C. Lamanna (2016). "Evidence of a derived titanosaurian (Dinosauria, Sauropoda) in the "Kem Kem beds" of Morocco, with comments on sauropod paleoecology in the Cretaceous of Africa". New Mexico Museum of Natural History and Science Bulletin. 71: 149–159.
36. Ute Richter; Alexander Mudroch; Lisa G. Buckley (2013). "Isolated theropod teeth from the Kem Kem Beds (Early Cenomanian) near Taouz, Morocco". Paläontologische Zeitschrift. 87 (2): 291–309. Bibcode:2013PalZ...87..291R. doi:10.1007/s12542-012-0153-1. S2CID 129382593.
37. Hendrickx, C.; Trapman, T. H.; Wills, S.; Holwerda, F. M.; Stein, K. H. W.; Rauhut, O. W. M.; Melzer, R. R.; Van Woensel, J.; Reumer, J. W. F. (2024). "A combined approach to identify isolated theropod teeth from the Cenomanian Kem Kem Group of Morocco: cladistic, discriminant, and machine learning analyses". Journal of Vertebrate Paleontology. 43 (4). e2311791. doi:10.1080/02724634.2024.2311791.
38. Robert S.H. Smyth; Nizar Ibrahim; Alexander Kao; David M. Martill (2019). "Abelisauroid cervical vertebrae from the Cretaceous Kem Kem beds of Southern Morocco and a review of Kem Kem abelisauroids". Cretaceous Research. 108: 104330. Bibcode:2020CrRes.10804330S. doi:10.1016/j.cretres.2019.104330. S2CID 214136033.
39. Brusatte, Stephen L.; Sereno, Paul C. (2007-12-12). "A new species of Carcharodontosaurus (Dinosauria: Theropoda) from the Cenomanian of Niger and a revision of the genus". Journal of Vertebrate Paleontology. 27 (4): 902–916. doi:10.1671/0272-4634(2007)27[902:ANSOCD]2.0.CO;2. ISSN 0272-4634. S2CID 86202969.
40. Paterna, Alessandro; Cau, Andrea (2022-10-11). "New giant theropod material from the Kem Kem Compound Assemblage (Morocco) with implications on the diversity of the mid-Cretaceous carcharodontosaurids from North Africa". Historical Biology. 35 (11): 2036–2044. doi:10.1080/08912963.2022.2131406. ISSN 0891-2963. S2CID 252856791.
41. "Table 4.1," in Weishampel, et al. (2004). Page 76.
42. Singer (2015). JURAPARK NA TROPIE NOWYCH DINOZAURÓW Z MAROKA (Online).
43. Molina-Pérez, Rubén; Larramendi, Asier; Connolly, David; Ramírez Cruz, Gonzalo Ángel; Mazzei, Sante; Atuchin, Andrey (2019-06-25). Dinosaur Facts and Figures: The Theropods and Other Dinosauriformes. Princeton University Press. doi:10.1515/9780691190594-013. ISBN 978-0-691-19059-4.
44. B. McFeeters (2013). "Bone "taxon" B: Reevaluation of a supposed small theropod dinosaur from the mid-Cretaceous of Morocco". Kirtlandia. 58: 38–41.
45. Andrea Cau; Fabio M. Dalla Vecchia; Matteo Fabbri (2012). "A thick-skulled theropod (Dinosauria, Saurischia) from the Upper Cretaceous of Morocco with implications for carcharodontosaurid cranial evolution". Cretaceous Research. 40: 251–260. doi:10.1016/j.cretres.2012.09.002.
46. Andrea Cau; Fabio Marco Dalla Vecchia; Matteo Fabbri (2012). "Evidence of a new carcharodontosaurid from the Upper Cretaceous of Morocco". Acta Palaeontologica Polonica. 57 (3): 661–665. doi:10.4202/app.2011.0043.
47. Ibrahim, Nizar; Maganuco, Simone; Dal Sasso, Cristiano; Fabbri, Matteo; Auditore, Marco; Bindellini, Gabriele; Martill, David M.; Zouhri, Samir; Mattarelli, Diego A.; Unwin, David M.; Wiemann, Jasmina (May 2020). "Tail-propelled aquatic locomotion in a theropod dinosaur". Nature. 581 (7806): 67–70. Bibcode:2020Natur.581...67I. doi:10.1038/s41586-020-2190-3. ISSN 1476-4687. PMID 32376955. S2CID 256820907.
48. David M. Martill; Roy Smith; David M. Unwin; Alexander Kao; James McPhee; Nizar Ibrahim (2020). "A new tapejarid (Pterosauria, Azhdarchoidea) from the mid-Cretaceous Kem Kem beds of Takmout, southern Morocco". Cretaceous Research. 112: Article 104424. Bibcode:2020CrRes.11204424M. doi:10.1016/j.cretres.2020.104424. S2CID 216303122.
49. Peter Wellnhofer; Eric Buffetaut (1999). "Pterosaur remains from the Cretaceous of Morocco". Paläontologische Zeitschrift. 73 (1–2): 133–142. Bibcode:1999PalZ...73..133W. doi:10.1007/BF02987987. S2CID 129032233.
50. Ibrahim, N.; Unwin, D.M.; Martill, D.M.; Baidder, L.; Zouhri, S. (2010). "A New Pterosaur (Pterodactyloidea: Azhdarchidae) from the Upper Cretaceous of Morocco". PLOS ONE. 5 (5): e10875. Bibcode:2010PLoSO...510875I. doi:10.1371/journal.pone.0010875. PMC 2877115. PMID 20520782.
51. Alexander Averianov (2014). "Review of taxonomy, geographic distribution, and paleoenvironments of Azhdarchidae (Pterosauria)". ZooKeys (432): 1–107. Bibcode:2014ZooK..432....1A. doi:10.3897/zookeys.432.7913. PMC 4141157. PMID 25152671.
52. Jacobs, Megan L.; Smith, Roy E.; Zouhri, Samir (2024). "A new ornithocheirid pterosaur (Pterosauria: Ornithocheiridae) from the mid-Cretaceous Ifezouane Formation, Kem Kem Group of Morocco". Cretaceous Research (in press): 106015. doi:10.1016/j.cretres.2024.106015.
53. Megan L. Jacobs; David M. Martill; David M. Unwin; Nizar Ibrahim; Samir Zouhri; Nicholas R. Longrich (2020). "New toothed pterosaurs (Pterosauria: Ornithocheiridae) from the middle Cretaceous Kem Kem beds of Morocco and implications for pterosaur palaeobiogeography and diversity". Cretaceous Research. 110: Article 104413. Bibcode:2020CrRes.11004413J. doi:10.1016/j.cretres.2020.104413. S2CID 214542129.
54. James McPhee; Nizar Ibrahim; Alex Kao; David M. Unwin; Roy Smith; David M. Martill (2020). "A new ?chaoyangopterid (Pterosauria: Pterodactyloidea) from the Cretaceous Kem Kem beds of Southern Morocco". Cretaceous Research. 110: Article 104410. Bibcode:2020CrRes.11004410M. doi:10.1016/j.cretres.2020.104410. S2CID 213739173.
55. Taissa Rodrigues; Alexander W. A. Kellner; Bryn J. Mader; Dale A. Russell (2011). "New pterosaur specimens from the Kem Kem beds (Upper Cretaceous, Cenomanian) of Morocco". Rivista Italiana di Paleontologia e Stratigrafia. 117 (1): 149–160. doi:10.13130/2039-4942/5967.
56. Borja Holgado; Rodrigo V. Pêgas (2020). "A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae". Acta Palaeontologica Polonica. 65 (4): 743–761. doi:10.4202/app.00751.2020. S2CID 222075296.
57. Roy E. Smith; David M. Martill; Alexander Kao; Samir Zouhri; Nicholas Longrich (2020). "A long-billed, possible probe-feeding pterosaur (Pterodactyloidea: ?Azhdarchoidea) from the mid-Cretaceous of Morocco, North Africa". Cretaceous Research. 118: Article 104643. doi:10.1016/j.cretres.2020.104643. S2CID 225201538.
58. Jacobs, Megan L.; Martill, David M.; Ibrahim, Nizar; Longrich, Nick (March 2019). "A new species of Coloborhynchus (Pterosauria, Ornithocheiridae) from the mid-Cretaceous of North Africa". Cretaceous Research. 95: 77–88. Bibcode:2019CrRes..95...77J. doi:10.1016/j.cretres.2018.10.018. ISSN 0195-6671. S2CID 134439172.
59. Rodrigues, Taissa; Kellner, Alexander W. A (2008). "Review of the pterodactyloid pterosaur Coloborhynchus" (PDF). Zitteliana. B 28: 219–228.