Kem Kem Beds

Kem Kem Beds Stratigraphic range: Cenomanian[1]
Type	Geological formation
Location
Country	Morocco  Algeria

The Kem Kem Beds (also referred to by various names including the Continental Red Beds and Continental intercalaire^[2]) is a geological formation along the border between Morocco and Algeria in southeastern Morocco, whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.^[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 Beds (specifically in regards 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

Crocodylomorphs

Crocodylomorphs reported from the Continental Red Beds
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Aegisuchus	Aegisuchus witmeri			"Partial braincase of a large individual with skull roof, temporal, and occipital regions."[4]	An aegyptosuchid.	Araripesuchus Elosuchus cherifiensis
Araripesuchus	Araripesuchus rattoides
Elosuchus	Elosuchus cherifiensis				An Elosuchid.
Hamadasuchus	Hamadasuchus rebouli				A Peirosaurid.
Kemkemia	K. auditorei	Errachidia Province, Morocco[5]		Known from an isolated caudal vertebra.[5]	Initially thought to be a neotheropod,[5] but subsequently discovered to be an indeterminate crocodyliform.[6]
Laganosuchus	Laganosuchus maghrebensisis				A Stomatosuchid.

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.

Dinosaurs reported from the Continental Red Beds
Genus	Species	Location	Stratigraphic position	Material	Notes	Images
Abelisauridae indet.[7]	Indeterminate			Isolated teeth.[7] Partial right femur.[8]	An indeterminate abelisaurid.	Carcharodontosaurus saharicus Deltadromeus agilis Spinosaurus aegypticus Majungasaurus
Carcharodontosaurus[1]	C. saharicus[1]	Ksar-es-Souk province, Morocco.[1]
Deltadromeus	D. agilis			"Partial skeleton, isolated limb elements."[9]
Dromaeosauridae indet.[7]	Indeterminate			Isolated teeth.[7]	An indeterminate dromaeosaurid.
cf. Elaphrosaurus	Indeterminate	Ksar-es-Souk province, Morocco.[1]			Fossils previously referred to cf. Elaphrosaurus are actually indeterminate theropod remains.
Inosaurus	I. tedreftensis			"Vertebrae."[10]
cf. Majungasaurus	Indeterminate	Ksar-es-Souk province, Morocco.[1]			Misidentified remains of an indeterminate abelisaurid.
Rebbachisaurus	R. garasbae	Ksar-es-Souk province, Morocco.[1]			A rebbachisaurid.
Saurischia indet.[11]	Indeterminate			Isolated vertebrae.[11]	An indeterminate saurischian.
Sauroniops[12]	S. pachytholus			"An isolated and almost complete left frontal."[13]	A carcharodontosaurid distinct from Carcharodontosaurus.[12][13]
Spinosaurus[1]	S. Maroccanus[1]	Ksar-es-Souk province, Morocco.[1]			A spinosaurid. Spinosaurus maroccanus

Pterosaurs

Pterosaurs of the Kem Kem Beds
Genus	Species	Location	Stratigraphic position	Abundance	Notes	Images
Alanqa[14]	A. saharica[14]					Alanqa saharica
Azhdarchidae indet.[15]	Indeterminate[15]			Two middle cervical vertebrae.[15]	Averianov (2014) considered it to be a specimen of Alanqa saharica.[16]
Azhdarchoidea indet.[15]	Indeterminate[15]			Humerus.[15]	Averianov (2014) considered it to be a specimen of Alanqa saharica.[16]
Dsungaripteroidea (sensu Kellner, 2003) indet.[15][17]	Indeterminate[15]			Anterior fragment from a jaw, possibly a mandible.[15]	Averianov (2014) considered it to be a specimen of Alanqa saharica.[16]
?Pteranodontidae indet.[14]	Indeterminate[14]			A fragment of bone, possibly the anterior section of rostrum.[14]	A possible pteranodontid[14] or a specimen of Alanqa saharica.[16]
Siroccopteryx[18]	S. moroccensis[18]				Classified by some authors as a species belonging to the genus Coloborhynchus.[14]
Tapejaridae indet.[19]	Indeterminate[19]			A fragment of bone originally interpreted as a fragment of anterior mandibular symphysis,[19] though it might pertain to the rostrum instead.[14]	Might belong to the family Thalassodromidae[14] or be a specimen of Alanqa saharica instead.[16]

Turtles

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

See also

• Aoufous Formation, within which lie 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. Michard, A. (2008). Continental evolution: the geology of Morocco : structure, stratigraphy, and tectonics of the Africa-Atlantic-Mediterranean Triple junction. Published by Springer, 2008. 424 pages. ISBN 3-540-77075-5, ISBN 978-3-540-77075-6
3. http://www3.imperial.ac.uk/newsandeventspggrp/imperialcollege/newssummary/news_29-2-2016-9-34-57
4. Casey M. Holliday and 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...7E0471H. doi:10.1371/journal.pone.0030471. PMC 3269432. PMID 22303441.
5. 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.
6. 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.
7. Ute Richter, Alexander Mudroch and 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.
8. Alfio Alessandro Chiarenza and 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.
9. "Table 4.1," in Weishampel, et al. (2004). Page 76.
10. "Table 4.1," in Weishampel, et al. (2004). Page 78.
11. B. McFeeters (2013). "Bone "taxon" B: Reevaluation of a supposed small theropod dinosaur from the mid-Cretaceous of Morocco". Kirtlandia 58: 38–41.
12. Andrea Cau, Fabio M. Dalla Vecchia and Matteo Fabbri (2012). "A thick-skulled theropod (Dinosauria, Saurischia) from the Upper Cretaceous of Morocco with implications for carcharodontosaurid cranial evolution". Cretaceous Research. in press. doi:10.1016/j.cretres.2012.09.002.
13. Andrea Cau, Fabio Marco Dalla Vecchia and 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.
14. 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.
15. 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.
16. Alexander Averianov (2014). "Review of taxonomy, geographic distribution, and paleoenvironments of Azhdarchidae (Pterosauria)". ZooKeys 432: 1–107. doi:10.3897/zookeys.432.7913.
17. I.e. a member of the clade containing the most recent common ancestor of Nyctosaurus and Quetzalcoatlus and all their descendants. See: Kellner, A.W.A., 2003. Pterosaur phylogeny and comments on the evolutionary history of the AN group. In: Buffetaut, E., Mazin, J.M. (Eds.), Evolution and Palaeobiology of Pterosaurs. Geological Society, London, Special Publication 217, 105–137
18. Rodrigues, Taissa; Kellner, Alexander W. A (2008). "Review of the pterodactyloid pterosaur Coloborhynchus" (PDF). Zitteliana B 28: 219–228.
19. Peter Wellnhofer, Eric Buffetaut (1999). "Pterosaur remains from the Cretaceous of Morocco". Paläontologische Zeitschrift 73 (1–2): 133–142. doi:10.1007/BF02987987.