Kem Kem Group | |
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Stratigraphic range: Cenomanian [1] ~ | |
Type | Geological group |
Sub-units | Douira Formation, Gara Sbaa Formation |
Underlies | Cenomanian-Turonian limestone platform (Akrabou Formation) |
Lithology | |
Primary | Sandstone |
Location | |
Coordinates | 32°50′N4°50′W / 32.833°N 4.833°W |
Approximate paleocoordinates | 18°48′N4°06′W / 18.8°N 4.1°W |
Region | Er Rachidia, Tafilalt |
Country | Morocco |
Extent | central and eastern Morocco north and south of the Pre-African Trough |
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.
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]
Cartilaginous fish | |||||
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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] | |||
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 | |||||
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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] | ||
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 | |||||
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Genus | Species | Location | Material | Notes | Images |
Arganodus | A. tiguidiensis | A lungfish | |||
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 | ||||||
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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 | ||||
Color key
| Notes Uncertain or tentative taxa are in small text; |
Lizards and snakes reported from the Continental Red Beds | ||||||
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Genus | Species | Location | Stratigraphic position | Material | Notes | Images |
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] | |||
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. | |||
N. begaa [24] | One posterior and two mid-trunk vertebrae | A stem-snake. | ||||
Indeterminate [24] | A mid-trunk vertebra | |||||
cf. S. libycus | Vertebrae [2] | An early snake. | ||||
Plesiosaurs | ||||||
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Genus | Species | Location | Stratigraphic position | Material | Notes | Images |
Leptocleididae cf. Leptocleidus [25] | indeterminate | |||||
Turtles reported from the Continental Red Beds | ||||||
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Genus | Species | Location | Stratigraphic position | Material | Notes | |
D. schaefferi | ||||||
G. emringeri | ||||||
G. whitei | ||||||
H. escuilliei | ||||||
A tooth enamel identified as cf. Sarcosuchus was discovered from the Ifezouane Formation. [26]
Crocodylomorphs reported from the Continental Red Beds | ||||||
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Genus | Species | Location | Stratigraphic position | Material | Notes | Images |
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] | ||||
A. taouzensis | Paired mandibles and a partial right mandible | A peirosaurid. | ||||
A. rattoides | Douira Formation | |||||
E. cherifiensis |
| An elosuchid. The material may represent two different species. [2] | ||||
H. rebouli |
| A peirosaurid. | ||||
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] | |||
L. thaumastos | ||||||
Lavocatchampsa sigogneaurusselae | Anterior portion of a rostrum with mandible, with an almost complete dentition [31] | |||||
Indeterminate lithostrotian remains once misattributed to the Titanosauridae are present in the province of Ksar-es-Souk, Morocco. [1]
Color key
| Notes Uncertain or tentative taxa are in small text; |
Ornithischians reported from the Continental Red Beds | ||||||
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Genus | Species | Location | Stratigraphic position | Material | Notes | Images |
Indeterminate | Douira Formation | An isolated tooth. [2] | A probable ankylosaur [32] | |||
Indeterminate | Douira Formation | A large, clover-shaped, three-toed footprint. [2] | Comparable in size and shape to tracks typically attributed to Iguanodon . [33] | |||
Sauropods reported from the Continental Red Beds | ||||||
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Genus | Species | Location | Stratigraphic position | Material | Notes | Images |
R. garasbae | Ksar-es-Souk province, Morocco. [1] | Gara Sbaa Formation | ||||
Indeterminate | 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] | |||||
Indeterminate |
| 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 reported from the Continental Red Beds | ||||||
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Genus | Species | Location | Stratigraphic position | Material | Notes | Images |
Indeterminate | Isolated teeth. [36] [37] | 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. [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 | |
D. agilis | Gara Sbaa Formation | "Partial skeleton, isolated limb elements." [41] | A noasaurid ceratosaurian or possible neovenatorid carnosaur. May be synonymous with Bahariasaurus . | |||
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. | |||
Indeterminate | ||||||
"Osteoporosia" [42] | "O. gigantea" [42] | A tooth and a possible neural arch from another specimen. [42] | A theropod, possibly synonymous with Sauroniops . [43] | |||
Indeterminate | An isolated cervical vertebra. [44] | An indeterminate saurischian. | ||||
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] | |||
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 of the Kem Kem Beds | ||||||
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Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
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] | ||
Akharhynchus [52] | A. martilli | Tafilalt | Ifezouane Formation | A fragment of the anterior part of the premaxillae | A tropeognathine anhanguerian. | |
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] | |||
A. cf. piscator [53] | upper Ifezouane Formation | Partial mandibular symphysis [53] | ||||
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] | 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] | ||
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] | ||
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. | |||
O. cf. simus. [53] | upper Ifezouane Formation | Premaxillae fragment [53] | ||||
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] |
Carcharodontosaurus is a genus of carnivorous theropod dinosaur that lived in North Africa from about 100 to 94 million years ago during the Cenomanian age of the Late Cretaceous. Two teeth of the genus, now lost, were first described from Algeria by French paleontologists Charles Depéret and Justin Savornin as Megalosaurus saharicus. A partial skeleton was collected by crews of German paleontologist Ernst Stromer during a 1914 expedition to Egypt. Stromer did not report the Egyptian find until 1931, in which he dubbed the novel genus Carcharodontosaurus, making the type species C. saharicus. Unfortunately, this skeleton was destroyed during the Second World War. In 1995 a nearly complete skull of C. saharicus, the first well-preserved specimen to be found in almost a century, was discovered in the Kem Kem Beds of Morocco; it was designated the neotype in 1996. Fossils unearthed from the Echkar Formation of northern Niger were described and named as another species, C. iguidensis, in 2007.
Sigilmassasaurus is a controversial genus of spinosaurid dinosaur that lived approximately 100 to 94 million years ago during the Late Cretaceous Period in what is now northern Africa. Named in 1996 by Canadian paleontologist Dale Russell, it contains a single species, Sigilmassasaurus brevicollis. The identity of the genus has been debated by scientists, with some considering its fossils to represent material from the closely related species Spinosaurus aegyptiacus, while others have classified it as a separate taxon, forming the clade Spinosaurini with Spinosaurus as its sister taxon.
Siroccopteryx is an extinct genus of anhanguerid pterodactyloid pterosaur which lived in Morocco during the Cenomanian stage of the Late Cretaceous. Some researchers, such as David M. Unwin, consider the genus a junior synonym of Coloborhynchus.
Coloborhynchus is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. Coloborhynchus is known from the Lower Cretaceous of England, and depending on which species are included, possibly the Albian and Cenomanian ages as well. Coloborhynchus was once thought to be the largest known toothed pterosaur, however, a specimen of the closely related Tropeognathus is now thought to have had a larger wingspan.
The Cambridge Greensand is a geological unit in England whose strata are earliest Cenomanian in age. It lies above the erosive contact between the Gault Formation and the Chalk Group in the vicinity of Cambridgeshire, and technically forms the lowest member bed of the West Melbury Marly Chalk Formation. It is a remanié deposit, containing reworked fossils of late Albian age, including those of dinosaurs and pterosaurs.
Bahariasauridae is a potential family of averostran theropods that might include a handful of African and South American genera, such as Aoniraptor, Bahariasaurus, Deltadromeus, and Gualicho. The placement of these theropods is controversial, with some studies placing them as basal ceratosaurs possibly related to Noasauridae, others classifying them as megaraptorans, basal neovenatorids, or basal coelurosaurs. There is also a possibility the group might not be monophyletic, as a monograph on the vertebrate diversity in the Kem Kem Beds published in 2020 found Bahariasaurus to be nomen dubium. In the same paper Deltadromeus is classified as an noasaurid, a result also recovered by some previous studies. A 2024 phylogenetic analysis found Aoniraptor, Bahariasaurus, Deltadromeus, and Gualicho to form a monophyletic clade as the sister taxon to Elaphrosaurus near the base of Ceratosauria.
Chaoyangopteridae is a family of pterosaurs within the larger group Azhdarchoidea. Chaoyangopterids lived mostly during the Early Cretaceous period, though possible members, Microtuban, Xericeps and Argentinadraco, may extend the fossil range to the Late Cretaceous.
Alanqa is a genus of pterodactyloid pterosaur from the Late Cretaceous period of what is now the Kem Kem Beds of southeastern Morocco. The name Alanqa comes from the Arabic word العنقاءal-‘Anqā’, for a mythical bird of Arabian culture.
Sauroniops is a controversial genus of carnivorous basal carcharodontosaurid theropod dinosaur known from the Late Cretaceous Gara Sbaa Formation, and possibly also the Kem Kem Formation, both of Morocco. The type, and currently only, species is S. platytholus.
Gualicho is a genus of theropod dinosaur. The type species is Gualicho shinyae. It lived in what is now northern Patagonia, on what was then a South American island continent split off from the supercontinent Gondwana. The fossils were found in the Huincul Formation, dating to the late Cenomanian-early Turonian age of the upper Cretaceous Period, around 91 million years ago.
Xericeps is a genus of pterosaur from the Cenomanian stage of the Late Cretaceous. It was discovered from the Kem Kem Beds of southeastern Morocco.
Nizar Ibrahim is a German-Moroccan vertebrate paleontologist and comparative anatomist. He is currently a senior lecturer at the University of Portsmouth. Ibrahim has led several expeditions to Africa's Sahara and is notable for his research on fossil vertebrates from the Kem Kem Group, including pterosaurs, crocodyliforms, and dinosaurs. In recent years, research led by Ibrahim radically changed ideas about the morphology and life habits of one of the largest predatory dinosaurs, Spinosaurus aegyptiacus. Ibrahim also has interests in bioinformatics and contributed to the NSF-funded Phenoscape project. He regularly engages with the public and is a speaker with the National Geographic Speakers Bureau.
Afrotapejara is an extinct genus of tapejarid pterosaur discovered in Morocco. The type species, Afrotapejara zouhri, was named and described in 2020. It was the first tapejarid discovered in Africa and the fourth pterosaur discovered in the Kem Kem Beds.
Apatorhamphus is an extinct genus of azhdarchoid pterosaur from the Kem Kem Group of Morocco. It might have been part of the Chaoyangopteridae. It is only known from a few snout fragments and it likely had a wingspan of between 3–7 metres (9.8–23.0 ft)
Leptostomia is a genus of long-beaked pterosaur from the mid-Cretaceous (Cenomanian) of Morocco, North Africa. The type species is L. begaaensis, which was named and described in 2021 from sediments of the Kem Kem Group in Morocco. It was a small animal with a long, slender bill which is thought to have been used to probe sediments for worms and other invertebrates, similar to kiwi birds and curlews. Leptostomia is likely a member of the Azhdarchoidea.
Nicorhynchus is a genus of anhanguerid pterosaur from the Cretaceous period. It contains two species, the type species, N. capito, from the Cambridge Greensand of England, and N. fluviferox from the Kem Kem Group of Morocco. These species were previously assigned to Coloborhynchus.
Oniichthys is an extinct genus of gar in the family Lepisosteidae. It contains a single species, O. falipoui, known from the Late Cretaceous (Cenomanian) of Morocco.
Akharhynchus is an extinct genus of tropeognathine pteranodontoid pterosaurs possibly from the Cretaceous Kem Kem Group of Morocco. The genus contains a single species, A. martilli, known from a small fragment of the premaxillae.