Kem Kem Group

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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
Lithology
PrimarySandstone
Location
RegionEr Rachidia, Tafilalt
Country Morocco
Extentcentral and eastern Morocco north and south of the Pre-African Trough
KemKem-fig1-map.jpg
(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]

Fish[edit]

Fishes
Genus Species Location Material Notes Images
Acrodontidae indet.[2] Indeterminate Members of Hybodontoidea
Life restoration of Aidachar
Life restoration of Axelrodichthys
Rostrum and teeth fossils from Onchopristis
The Queensland Lungfish, the only living member of Neoceratodus
Adrianaichthys[2] A. pankowskii Isolated scales[4] and two skulls[5] A member of Lepisosteiformes. Originally described as a species of Lepidotes, but subsequently transferred to a separate genus.[6]
Agassizilia[7] 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[8] or Ophiopsiellidae.[9]
Aidachar A. pankowskii
Arganodus A. tiguidiensis
Axelrodichthys[10] A.? lavocati A mawsoniid coelacanth
Bahariyodon[2] B. bartheli A member of Hybodontoidea
Bartschichthys[2] Bartschichthys sp. Isolated pinnulae (spines that support each dorsal finlet)[2] A cladistian
Bawitius cf. Bawitius sp. Isolated scales and jaw fragments[4]
Calamopleurus[2] C. africanus A partial skull[2] A member of Amiiformes
Cenocarcharias[2] C. tenuiplicatus One tooth[2] A member of the family Cretoxyrhinidae
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
Distobatus[2] D. nutiae A member of Hybodontoidea
Erfoudichthys[2] E. rosae Isolated skull[2] A small-bodied teleost of unknown affinity
Haimirichia[2] H. amonensis One tooth[2] A mackerel shark
Marckgrafia[2] M. lybica 13 teeth[2] A member of Batoidea
Neoceratodus N. africanus
Neoproscinetes[7] N. africanus A member of the family Pycnodontidae
Obaichthys O. africanus Isolated scales[4] A member of Lepisosteiformes
Onchopristis O. numidus A giant sawfish[11]
Oniichthys O. falipoui Near complete skeleton including skull[4] A member of Lepisosteiformes
Palaeonotopterus[2] P. greenwoodi A member of Osteoglossomorpha
Peyeria[2] P. libyca Three teeth[2] A sawfish. Might be a junior synonym of Onchopristis numidus.
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
Tribodus[2] Tribodus sp. A member of Hybodontoidea

Amphibians[edit]

Amphibians
Genus Species Location Stratigraphic position Material Notes Images
Anura indet.[12] Indeterminate Partial braincase, jaw fragments and procoelous vertebrae[2] Fossil material probably pertaining to several species of non-pipid frogs.[2]
cf. Kababisha[12] Indeterminate A salamander belonging to the family Sirenidae
Oumtkoutia[12] O. anae A frog belonging to the family Pipidae

Lizards and snakes[edit]

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

Bicuspidon

Bicuspidon hogreli[13]

A polyglyphanodontid lizard.

Jeddaherdan[14]

Jeddaherdan aleadonta

Partial mandible with teeth.

An iguanian belonging to the group Acrodonta, possibly a relative of the uromastycine agamids.

Lapparentophis

Lapparentophis ragei[15]

Two isolated trunk vertebrae

An early snake.

Madtsoiidae indet.[12]

Indeterminate

Vertebrae[2]

An early snake.

?Nigerophiidae indet.[12]

Indeterminate

Dorsal vertebrae[2]

An early snake.

Norisophis[16]

Norisophis begaa[16]

One posterior and two mid-trunk vertebrae

A stem-snake.

Indeterminate[16]

A mid-trunk vertebra

Simoliophis[12]

cf. Simoliophis libycus

Vertebrae[2]

An early snake.

Crocodylomorphs[edit]

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."[17]

An aegyptosuchid.

Araripesuchus

Araripesuchus rattoides

Elosuchus

Elosuchus cherifiensis

An Elosuchid.

Hamadasuchus

Hamadasuchus rebouli

A Peirosaurid.

Kemkemia

K. auditorei

Errachidia Province, Morocco[18]

Known from an isolated caudal vertebra.[18]

Initially thought to be a neotheropod,[18] but subsequently discovered to be an indeterminate crocodyliform.[19]

Laganosuchus

Laganosuchus maghrebensisis

A Stomatosuchid.

Lavocatchampsa[20]

Lavocatchampsa sigogneaurusselae

Anterior portion of a rostrum with mandible, with an almost complete dentition[20]

A candidodontid notosuchian.[20]

Dinosaurs[edit]

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[21]

Indeterminate

Isolated teeth.[21]
Partial right femur.[22] Partial right maxilla.[2] A cervical vertebra.[23] Left maxilla.[2]

A large indeterminate abelisaurid closely related to Rugops.[2]

Carcharodontosaurus[1]

C. saharicus[1]

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

A carcharodontosaurid theropod.

Deltadromeus

D. agilis

Gara Sbaa Formation

"Partial skeleton, isolated limb elements."[24]

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

Dromaeosauridae[21]

Indeterminate

Isolated teeth.[21]

An indeterminate dromaeosaurid.

cf. Elaphrosaurus

Indeterminate

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

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

Noasauridae[23]

Indeterminate

An anterior cervical vertebra[23]

Ornithischia[2]

Indeterminate

An isolated tooth.[2]

A small-bodied ornithischian of uncertain phylogenetic placement, possibly a thyreophoran[2]

Ornithopoda[25]

Indeterminate

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

Comparable in size and shape to tracks typically attributed to Iguanodon.[25]

Rebbachisaurus

R. garasbae

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

Gara Sbaa Formation

A rebbachisaurid.

Saurischia[26]

Indeterminate

An isolated cervical vertebra.[26]

An indeterminate saurischian.

Sauroniops[27]

S. pachytholus

"An isolated and almost complete left frontal."[28]

A carcharodontosaurid distinct from Carcharodontosaurus.[27][28]

Somphospondyli[29]

Indeterminate

Anterior dorsal vertebra, partial right ischium[29]

The vertebra might belong to a basal titanosaurian, possibly distinct from Aegyptosaurus and Paralititan.[29] 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.[29]

Spinosaurus

S. aegyptiacus

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

Titanosauria[30][2]

Indeterminate

  • 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]

Rugops R. primus Ksar-es-Souk, Morroco Douria Formation Left maxillae.

Pterosaurs[edit]

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

Afrotapejara[31]

A. zouhri[31]

A fragment of bone interpreted as a fragment of anterior mandibular symphysis,[32] and additional jaw fragments that pertain to the rostrum.[33]

A tapejarid pterosaur. Originally believed to belong to either the family Thalassodromidae[33] or an additional specimen of Alanqa saharica.[34]

Alanqa[33]

A. saharica[33]

Anhanguera[35]

A. cf. piscator[35]

Partial mandibular symphysis[35]

Apatorhamphus[36]

A. gyrostega[36]

Partial rostrum and mandible[36]

A possible chaoyangopterid azhdarchoid pterosaur.[36] Originally believed to be a possible pteranodontid,[33] a possible dsungaripterid,[37] a possible non-azhdarchid azhdarchoid or nyctosaurid,[37] or a specimen of Alanqa saharica.[34]

Azhdarchidae indet.[37]

Indeterminate[37]

Three middle cervical vertebrae.[37][33]

Averianov (2014) considered these vertebrae to pertain to Alanqa saharica,[34] although the vertebrae may be indicative of two taxa.[37]

Coloborhynchus[35] C. sp. A.[35] Hassi El Begaa Premaxillae fragment[35] Possibly a specimen of Nicorhynchus fluviferox.[38]

Leptostomia[39]

L. begaaensis[39]

Aferdou N’ Chaft

Partial rostrum and partial mandibular synthesis[39]

A small, long-beaked pterosaur, likely a member of Azhdarchoidea.[39]

Nicorhynchus[38]

N. fluviferox[40][38]

Possibly Aferdou N’Chaft, Hassi El Begaa[38]

An anterior portion of the rostrum.[38]

Originally described as a species of Coloborhynchus[40] but subsequently transferred to the genus Nicorhynchus.

Ornithocheirus[35]

O. cf. simus.[35]

Premaxillae fragment[35]

Siroccopteryx[41]

S. moroccensis[41]

Classified by some authors as a species belonging to the genus Coloborhynchus.[33]

Xericeps X. curvirostra A mandible fragment

Turtles[edit]

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

Dirqadim

Dirqadim schaefferi

A Euraxemydid

Galianemys

G. emringeri

A Cearachelyin

G. whitei

Hamadachelys

Hamadachelys escuilliei

See also[edit]

References[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 av 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. ^ 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.
  5. ^ https://palaeo-electronica.org/2011_1/239/239.pdf
  6. ^ 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.
  7. ^ 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.
  8. ^ 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.
  9. ^ 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.
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