Kem Kem Beds

Kem Kem Talis
Stratigraphic range: Cenomanian^[1]
Kem Kem Talis; 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 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]

Restorations of life in the Kem Kem beds

Vertebrate paleofauna

Fishes

Fishes reported from the Continental Red Beds
Genus	Species	Location	Material	Notes	Images
Aidachar	Aidachar pankowskii
Arganodus	Arganodus tiguidiensis				Rostrum and teeth fossils from Onchopristis Life restoration of Mawsonia
Bawitius	cf. Bawitius sp.		Isolated scales and jaw fragments^[4]
Lepidotes	Lepidotes pankowskii		Isolated scales^[4] and two skulls^[5]
Mawsonia	Mawsonia lavocati
Neoceratodus	Neoceratodus africanus
Obaichthys	Obaichthys africanus		Isolated scales^[4]
Onchopristis	Onchopristis numidus			A giant sawfish^[6]
Oniichthys	Oniichthys falipoui		Near complete skeleton including skull^[4]
Stromerichthys	Stromerichthys aethiopicus

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."^[7]	An aegyptosuchid.	Araripesuchus Elosuchus cherifiensis
Araripesuchus	Araripesuchus rattoides
Elosuchus	Elosuchus cherifiensis				An Elosuchid.
Hamadasuchus	Hamadasuchus rebouli				A Peirosaurid.
Kemkemia	K. auditorei	Errachidia Province, Morocco^[8]		Known from an isolated caudal vertebra.^[8]	Initially thought to be a neotheropod,^[8] but subsequently discovered to be an indeterminate crocodyliform.^[9]
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.^[10]	Indeterminate			Isolated teeth.^[10] Partial right femur.^[11]	An indeterminate abelisaurid.	Carcharodontosaurus saharicus Rebbachisaurus Spinosaurus aegypticus reconstruction
Carcharodontosaurus^[1]	C. saharicus^[1]	Ksar-es-Souk province, Morocco.^[1]			A carcharodontosaurid theropod.
Deltadromeus	D. agilis			"Partial skeleton, isolated limb elements."^[12]	A noasaurid ceratosaurian or possible neovenatorid carnosaur. May be synonymous with Bahariasaurus.
Dromaeosauridae indet.^[10]	Indeterminate			Isolated teeth.^[10]	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."^[13]
Rebbachisaurus	R. garasbae	Ksar-es-Souk province, Morocco.^[1]			A rebbachisaurid.
cf. Brachiosaurus	"B. nougaredi"	Algeria		Partial sacrum, phalanges	Fossils assigned to Brachiosaurus are actually from an indeterminate Titanosauriform, still assigned to "B. nougaredi".^[14]
Saurischia indet.^[15]	Indeterminate			Isolated vertebrae.^[15]	An indeterminate saurischian.
Sauroniops^[16]	S. pachytholus			"An isolated and almost complete left frontal."^[17]	A carcharodontosaurid distinct from Carcharodontosaurus.^[16]^[17]
Sigilmassasaurus	S. brevicollis	Ksar-es-Souk province, Morocco.
Spinosaurus	S. maroccanus	Ksar-es-Souk province, Morocco.^[1]

Pterosaurs

Pterosaurs of the Kem Kem Beds
Genus	Species	Location	Stratigraphic position	Abundance	Notes	Images
Alanqa^[18]	A. saharica^[18]					Alanqa saharica
Azhdarchidae indet.^[19]	Indeterminate^[19]			Three middle cervical vertebrae.^[19]^[18]	Averianov (2014) considered these vertebrae to pertain to Alanqa saharica,^[20] although the vertebrae may be indicative of two taxa.^[19]
Azhdarchoidea indet.^[19]	Indeterminate^[19]			Humerus.^[19]	Averianov (2014) considered it to be a specimen of Alanqa saharica.^[20]
Dsungaripteroidea (sensu Kellner, 2003) indet.^[19]^[21]	Indeterminate^[19]			Anterior fragment from a jaw, possibly a mandible.^[19]	May be a specimen of Alanqa saharica^[20] or a non-azhdarchid azhdarchoid or nyctosaurid.^[19]
?Pteranodontidae indet.^[18]	Indeterminate^[18]			A fragment of bone, possibly the anterior section of rostrum.^[18]	A possible pteranodontid^[18] or a specimen of Alanqa saharica.^[20]
Siroccopteryx^[22]	S. moroccensis^[22]				Classified by some authors as a species belonging to the genus Coloborhynchus.^[18]
Tapejaridae indet.^[23]	Indeterminate^[23]			A fragment of bone originally interpreted as a fragment of anterior mandibular symphysis,^[23] though it might pertain to the rostrum instead.^[18]	Might belong to the family Thalassodromidae^[18] or be a specimen of Alanqa saharica instead.^[20]
Xericeps	X. curvirostra			A mandible fragment

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

References

1 2 3 4 5 6 7 8 9 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.
↑ 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
↑ http://www3.imperial.ac.uk/newsandeventspggrp/imperialcollege/newssummary/news_29-2-2016-9-34-57
1 2 3 4 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. doi:10.1371/journal.pone.0125786. ISSN 1932-6203. PMC 4446216. PMID 26018561.
↑ https://palaeo-electronica.org/2011_1/239/239.pdf
↑ "Onchopristis - Paleobiology Database - Details - Encyclopedia of Life". Encyclopedia of Life. Retrieved 2018-07-07.
↑ 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...7E0471H. doi:10.1371/journal.pone.0030471. PMC 3269432. PMID 22303441.
1 2 3 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.
↑ 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.
1 2 3 4 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.
↑ 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.
↑ "Table 4.1," in Weishampel, et al. (2004). Page 76.
↑ "Table 4.1," in Weishampel, et al. (2004). Page 78.
↑ Upchurch, P., Barrett, P.M. & Dodson, P. (2004). "Sauropoda", pp. 259–322 in Weishampel, D.B., Dodson, P. & Osmólska, H. (Eds.) The Dinosauria, 2nd edition. University of California Press, Berkeley and Los Angeles. ISBN 0520254082.
1 2 B. McFeeters (2013). "Bone "taxon" B: Reevaluation of a supposed small theropod dinosaur from the mid-Cretaceous of Morocco". Kirtlandia. 58: 38–41.
1 2 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. in press: 251–260. doi:10.1016/j.cretres.2012.09.002.
1 2 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. Archived from the original on 2012-11-08.
1 2 3 4 5 6 7 8 9 10 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.
1 2 3 4 5 6 7 8 9 10 11 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.
1 2 3 4 5 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.
↑ 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
1 2 Rodrigues, Taissa; Kellner, Alexander W. A (2008). "Review of the pterodactyloid pterosaur Coloborhynchus" (PDF). Zitteliana. B 28: 219–228.
1 2 3 Peter Wellnhofer; Eric Buffetaut (1999). "Pterosaur remains from the Cretaceous of Morocco". Paläontologische Zeitschrift. 73 (1–2): 133–142. doi:10.1007/BF02987987.

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[cretaceousdistribution-1] 1 2 3 4 5 6 7 8 9 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.

[continental-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] ttp://www3.imperial.ac.uk/newsandeventspggrp/imperialcollege/newssummary/news_29-2-2016-9-34-57

[Cavin-4] 1 2 3 4 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. doi:10.1371/journal.pone.0125786. ISSN 1932-6203. PMC 4446216. PMID 26018561.

[5] ttps://palaeo-electronica.org/2011_1/239/239.pdf

[6] "Onchopristis - Paleobiology Database - Details - Encyclopedia of Life". Encyclopedia of Life. Retrieved 2018-07-07.

[HG12-7] 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...7E0471H. doi:10.1371/journal.pone.0030471. PMC 3269432. PMID 22303441.

[kemkemia-8] 1 2 3 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.

[9] 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.

[RichteretalPZ-10] 1 2 3 4 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.

[11] 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.

[table-4-1-76-12] "Table 4.1," in Weishampel, et al. (2004). Page 76.

[table-4-1-78-13] "Table 4.1," in Weishampel, et al. (2004). Page 78.

[Up-14] Upchurch, P., Barrett, P.M. & Dodson, P. (2004). "Sauropoda", pp. 259–322 in Weishampel, D.B., Dodson, P. & Osmólska, H. (Eds.) The Dinosauria, 2nd edition. University of California Press, Berkeley and Los Angeles. ISBN 0520254082.

[McFeeters-15] 1 2 B. McFeeters (2013). "Bone "taxon" B: Reevaluation of a supposed small theropod dinosaur from the mid-Cretaceous of Morocco". Kirtlandia. 58: 38–41.

[Sauroniops-16] 1 2 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. in press: 251–260. doi:10.1016/j.cretres.2012.09.002.

[CauetalAPP-17] 1 2 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. Archived from the original on 2012-11-08.

[ibrahimetal2010-18] 1 2 3 4 5 6 7 8 9 10 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.

[RIPS2011-19] 1 2 3 4 5 6 7 8 9 10 11 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.

[ZK2014-20] 1 2 3 4 5 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.

[21] 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

[Coloborhynchus-22] 1 2 Rodrigues, Taissa; Kellner, Alexander W. A (2008). "Review of the pterodactyloid pterosaur Coloborhynchus" (PDF). Zitteliana. B 28: 219–228.

[PZ1999-23] 1 2 3 Peter Wellnhofer; Eric Buffetaut (1999). "Pterosaur remains from the Cretaceous of Morocco". Paläontologische Zeitschrift. 73 (1–2): 133–142. doi:10.1007/BF02987987.