Kem Kem Group

From Wikipedia, the free encyclopedia
(Redirected from Kem Kem Beds)
Kem Kem Group
Stratigraphic range: Cenomanian[1]
~98–92.5 Ma
TypeGeological group
Unit ofHamadian Supergroup
Sub-unitsDouira Formation, Gara Sbaa Formation
UnderliesCenomanian-Turonian limestone platform
OverliesUnconformity with Paleozoic rocks
RegionEr Rachidia, Tafilalt
Country Morocco
Extentcentral and eastern Morocco north and south of the Pre-African Trough
(A) Location of the Kem Kem Group; (B) Coeval sites in North Africa; (C) Key Cretaceous-aged outcrops

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, the lower Gara Sbaa Formation and the upper Douira Formation.[2] It is exposed on an escarpment along the Algeria–Morocco border.

The unit unconformably overlies Paleozoic marine units of Cambrian, Silurian and Devonian age, and is itself capped by limestone platform rock of Cenomanian-Turonian age. It primarily consists of deltaic deposits. The lower Gara Sbaa Formation primarily consists of fine and medium grained sandstone, while the Douira Formation consists of fining-upwards, 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 be separated by millions of years.[3]

Vertebrate paleofauna[edit]

Cartilaginous fish[edit]

Cartilaginous fish
Genus Species Location Material Notes Images
Acrodontidae indet.[2] Indeterminate Members of Hybodontoidea
Rostrum and teeth fossils from Onchopristis
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. numidus A rajiform ray[4]
Peyeria[2] P. libyca Three teeth[2] A sawfish. Might be a junior synonym of Onchopristis numidus.
Tribodus[2] Tribodus sp. A member of Hybodontoidea

Ray-finned fish[edit]

Ray-finned fish
Genus Species Location Material Notes Images
Adrianaichthys[2] A. pankowskii Isolated scales[5] and two skulls[6] A member of Lepisosteiformes. Originally described as a species of Lepidotes, but subsequently transferred to a separate genus.[7]
Life restoration of Aidachar
Agassizilia[8] 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[9] or Ophiopsiellidae.[10]
Aidachar A. pankowskii A member of Ichthyodectiformes
Bartschichthys[2] Bartschichthys sp. Isolated pinnulae (spines that support each dorsal finlet)[2] A cladistian
Bawitius cf. Bawitius sp. Isolated scales and jaw fragments[5] 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] Diplomystus 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[8] N. africanus A member of the family Pycnodontidae
Obaichthys O. africanus Isolated scales[5] A member of Lepisosteiformes
Oniichthys O. falipoui Near complete skeleton including skull[5] 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] Sudania sp. An isolated pinnula[2] A cladistian

Lobe-finned fish[edit]

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[11] A.? lavocati A mawsoniid coelacanth
Neoceratodus N. africanus A lungfish


Genus Species Location Stratigraphic position Material Notes Images
Anura indet.[12] Indeterminate Douira Formation Incomplete left ilium
Cretadhefdaa[12] 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[13] Indeterminate A salamander belonging to the family Sirenidae
?Neobatrachia indet.[12] Indeterminate Douira Formation Incomplete humerus A frog, possibly a member of Ranoidea.
Oumtkoutia[13] O. anae A frog belonging to the family Pipidae

Lizards and snakes[edit]

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.
Lizards and snakes reported from the Continental Red Beds
Genus Species Location Stratigraphic position Material Notes Images


Bicuspidon hogreli[14]

A polyglyphanodontid lizard.


Jeddaherdan 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.[16]


Lapparentophis ragei[17]

Two isolated trunk vertebrae

An early snake.

Madtsoiidae indet.[13]



An early snake.

?Nigerophiidae indet.[13]


Dorsal vertebrae[2]

An early snake.


Norisophis begaa[18]

One posterior and two mid-trunk vertebrae

A stem-snake.


A mid-trunk vertebra


cf. Simoliophis libycus


An early snake.


Genus Species Location Stratigraphic position Material Notes Images
Leptocleididae indet.[19] Indeterminate


Turtles reported from the Continental Red Beds
Genus Species Location Stratigraphic position Material Notes


Dirqadim schaefferi

A Euraxemydid


G. emringeri

A Cearachelyin

G. whitei


Hamadachelys escuilliei


Crocodylomorphs reported from the Continental Red Beds
Genus Species Location Stratigraphic position Material Notes Images


Aegisuchus witmeri

"Partial braincase of a large individual with skull roof, temporal, and occipital regions."[20]

An aegyptosuchid that may be a synonym of Laganosuchus.[2]


Antaeusuchus taouzensis

Paired mandibles and a partial right mandible

A peirosaurid.


Araripesuchus rattoides

Douira Formation


Elosuchus cherifiensis

  • Gara Sbaa Formation
  • Douira Formation

An Elosuchid. The material may represent two different species.[2]


Hamadasuchus rebouli

  • Gara Sbaa Formation
  • Douira Formation?

A Peirosaurid.


K. auditorei

Errachidia Province, Morocco[22]

Known from an isolated caudal vertebra.[22]

Initially thought to be a neotheropod,[22] but subsequently discovered to be an indeterminate crocodyliform.[23]


Laganosuchus maghrebensisis

A Stomatosuchid.


Lavocatchampsa sigogneaurusselae

Anterior portion of a rostrum with mandible, with an almost complete dentition[24]

A candidodontid notosuchian.[24]


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
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.
Dinosaurs reported from the Continental Red Beds
Genus Species Location Stratigraphic position Material Notes Images



Isolated teeth.[25]
Partial right femur.[26] Partial right maxilla.[2] A cervical vertebra.[27] Left maxilla.[2]

Abelisaurid material belonging to one or two distinct taxa.[2]


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.

(Assigned as neotype for C. saharicus)[28]

A carcharodontosaurid theropod.

Carcharodontosauridae[29] Indeterminate Southeast of Taouz, Errachidia Province Gara Sbaa Formation partial maxilla and partial jugal A carcharodontosaurid theropod different from C. saharicus


D. agilis

Gara Sbaa Formation

"Partial skeleton, isolated limb elements."[30]

A noasaurid ceratosaurian or possible neovenatorid carnosaur. May be synonymous with Bahariasaurus.



Isolated teeth.[25]

An indeterminate dromaeosaurid.

cf. Elaphrosaurus


Ksar-es-Souk province, Morocco.[1]

Fossils previously referred to cf. Elaphrosaurus are actually indeterminate theropod remains.



An anterior cervical vertebra[27]



Douira Formation

An isolated tooth.[2]

A probable ankylosaur[31]



Douira Formation

A large, clover-shaped, three-toed footprint.[2]

Comparable in size and shape to tracks typically attributed to Iguanodon.[32]


R. garasbae

Ksar-es-Souk province, Morocco.[1]

Gara Sbaa Formation

A rebbachisaurid.



An isolated cervical vertebra.[33]

An indeterminate saurischian.


S. pachytholus

Gara Sbaa Formation

"An isolated and almost complete left frontal."[35]

A carcharodontosaurid distinct from Carcharodontosaurus.[34][35]



Anterior dorsal vertebra, partial right ischium[36]

The vertebra might belong to a basal titanosaurian, possibly distinct from Aegyptosaurus and Paralititan.[36] 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.[36]


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. (Assigned as neotype for S. aegyptiacus.)[37]

Numerous isolated bones.



  • Gara Sbaa Formation
  • Douira Formation

Isolated teeth, caudal vertebrae, a partial humerus, a tarsal bone and the proximal end of an ulna.[2]

Fossil material pertaining to one or more titanosaurian sauropods. Some fossils are indicative of large body size comparable to Paralititan stromeri.[2]


Pterosaurs of the Kem Kem Beds
Genus Species Location Stratigraphic position Abundance Notes Images


A. zouhri[39]

A fragment of bone interpreted as a fragment of anterior mandibular symphysis,[40] and additional jaw fragments that pertain to the rostrum.[41]

A tapejarid pterosaur. Originally believed to belong to either the family Thalassodromidae[41] or an additional specimen of Alanqa saharica.[42]


A. saharica[41]


A. cf. piscator[43]

Partial mandibular symphysis[43]


A. gyrostega[44]

Gara Sbaa Formation

Partial rostrum and mandible[44]

A possible chaoyangopterid azhdarchoid pterosaur.[44] Originally believed to be a possible pteranodontid,[41] a possible dsungaripterid,[45] a possible non-azhdarchid azhdarchoid or nyctosaurid,[45] or a specimen of Alanqa saharica.[42]

Azhdarchidae indet.[45]


Three middle cervical vertebrae.[45][41]

Averianov (2014) considered these vertebrae to pertain to Alanqa saharica,[42] although the vertebrae may be indicative of two taxa.[45]

Coloborhynchus[43] C. sp. A.[43] Hassi El Begaa Premaxillae fragment[43] Possibly a specimen of Nicorhynchus fluviferox.[46]


L. begaaensis[47]

Aferdou N’ Chaft

Gara Sbaa Formation

Partial rostrum and partial mandibular synthesis[47]

A small, long-beaked pterosaur, likely a member of Azhdarchoidea.[47]


N. fluviferox[48][46]

Possibly Aferdou N’Chaft, Hassi El Begaa[46]

Gara Sbaa Formation

An anterior portion of the rostrum.[46]

Originally described as a species of Coloborhynchus[48] but subsequently transferred to the genus Nicorhynchus.


O. cf. simus.[43]

Gara Sbaa Formation

Premaxillae fragment[43]


S. moroccensis[49]

Classified by some authors as a species belonging to the genus Coloborhynchus.[41]

Xericeps X. curvirostra Douira Formation A mandible fragment
X. sp. Gara Sbaa Formation Originally assigned to Alanqa, but is similar to Xericeps.[50]

See also[edit]


  1. ^ a b c d e f g h i 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. ^ 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 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. doi:10.3897/zookeys.928.47517. PMC 7188693. PMID 32362741.
  3. ^ Smith, Collin (2016-02-29). "Fossil find reveals just how big carnivorous dinosaur may have grown". Imperial News. Retrieved 2020-02-03.
  4. ^ 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.
  5. ^ a b c d 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.
  6. ^[bare URL PDF]
  7. ^ 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.
  8. ^ a b 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.
  9. ^ 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. doi:10.1111/j.1475-4983.2009.00851.x.
  10. ^ 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. doi:10.1080/02724634.2017.1420071. S2CID 89886438.
  11. ^ 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.
  12. ^ a b c 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.
  13. ^ a b c d e 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. doi:10.1127/pala/285/2008/1.
  14. ^ 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.
  15. ^ 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.
  16. ^ 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. 105412. doi:10.1016/j.cretres.2022.105412.
  17. ^ 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. doi:10.1016/j.crpv.2019.08.004.
  18. ^ a b c 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. doi:10.1016/j.cretres.2016.12.001.
  19. ^ 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. doi:10.1016/j.cretres.2022.105310.
  20. ^ 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.
  21. ^ 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. doi:10.1098/rsos.211254. PMID 34659786.
  22. ^ a b c 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.
  23. ^ 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.
  24. ^ a b c 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. doi:10.1080/02724634.2016.1211534. S2CID 89354184.
  25. ^ a b c d 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. doi:10.1007/s12542-012-0153-1. S2CID 129382593.
  26. ^ 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.
  27. ^ a b c 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. doi:10.1016/j.cretres.2019.104330.
  28. ^ 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.
  29. ^ 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: 1–9. doi:10.1080/08912963.2022.2131406. ISSN 0891-2963.
  30. ^ "Table 4.1," in Weishampel, et al. (2004). Page 76.
  31. ^ a b 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. doi:10.1016/j.cretres.2021.104881. ISSN 0195-6671.
  32. ^ a b 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.
  33. ^ a b B. McFeeters (2013). "Bone "taxon" B: Reevaluation of a supposed small theropod dinosaur from the mid-Cretaceous of Morocco". Kirtlandia. 58: 38–41.
  34. ^ a b 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.
  35. ^ a b 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.
  36. ^ a b c d 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.
  37. ^ 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. doi:10.1038/s41586-020-2190-3. ISSN 1476-4687.
  38. ^ 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.
  39. ^ a b 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. doi:10.1016/j.cretres.2020.104424.
  40. ^ Peter Wellnhofer; Eric Buffetaut (1999). "Pterosaur remains from the Cretaceous of Morocco". Paläontologische Zeitschrift. 73 (1–2): 133–142. doi:10.1007/BF02987987. S2CID 129032233.
  41. ^ a b c d e f g 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.
  42. ^ a b c Alexander Averianov (2014). "Review of taxonomy, geographic distribution, and paleoenvironments of Azhdarchidae (Pterosauria)". ZooKeys (432): 1–107. doi:10.3897/zookeys.432.7913. PMC 4141157. PMID 25152671.
  43. ^ a b c d e f g h i 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. doi:10.1016/j.cretres.2020.104413.
  44. ^ a b c d 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. doi:10.1016/j.cretres.2020.104410.
  45. ^ a b c d e f 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.
  46. ^ a b c d e 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.
  47. ^ a b c d 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.
  48. ^ a b 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. doi:10.1016/j.cretres.2018.10.018. ISSN 0195-6671.
  49. ^ a b Rodrigues, Taissa; Kellner, Alexander W. A (2008). "Review of the pterodactyloid pterosaur Coloborhynchus" (PDF). Zitteliana. B 28: 219–228.
  50. ^ Pêgas, Rodrigo V.; Holgado, Borja; Ortiz David, Leonardo D.; Baiano, Mattia A.; Costa, Fabiana R. (January 2022). "On the pterosaur Aerotitan sudamericanus (Neuquén Basin, Upper Cretaceous of Argentina), with comments on azhdarchoid phylogeny and jaw anatomy". Cretaceous Research. 129: 104998. doi:10.1016/j.cretres.2021.104998. ISSN 0195-6671.