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Kem Kem Group

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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.

Kem Kem Group
Stratigraphic range: Cenomanian[1]
~98–94 Ma
↓
TypeGeological group
Sub-unitsDouira Formation, Gara Sbaa Formation
UnderliesCenomanian-Turonian limestone platform (Akrabou Formation)
Lithology
PrimarySandstone
Location
Coordinates32°50′N 4°50′W / 32.833°N 4.833°W / 32.833; -4.833
Approximate paleocoordinates18°48′N 4°06′W / 18.8°N 4.1°W / 18.8; -4.1
RegionEr Rachidia, Tafilalt
Country Morocco
Extentcentral and eastern Morocco north and south of the Pre-African Trough
Kem Kem Group is located in Morocco
Kem Kem Group
Kem Kem Group (Morocco)

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]

Vertebrate paleofauna

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Cartilaginous fish

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Cartilaginous fish
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]
 
Rostrum and teeth fossils from Onchopristis
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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Ray-finned fish
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]
 
Life restoration of Aidachar
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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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[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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Amphibians
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

Lizards and snakes

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

Bicuspidon

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]

Lapparentophis

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.

Norisophis[24]

N. begaa[24]

One posterior and two mid-trunk vertebrae

A stem-snake.

Indeterminate[24]

A mid-trunk vertebra

Simoliophis[19]

cf. S. libycus

Vertebrae[2]

An early snake.

Plesiosaurs

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Plesiosaurs
Genus Species Location Stratigraphic position Material Notes Images
Leptocleididae cf. Leptocleidus[25] indeterminate

Turtles

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

Dirqadim

D. schaefferi

A Euraxemydid

Galianemys

G. emringeri

A Cearachelyin

G. whitei

Hamadachelys

H. escuilliei

Crocodylomorphs

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A tooth enamel identified as cf. Sarcosuchus was discovered from the Ifezouane Formation.[26]

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

Aegisuchus

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]

 
Aegisuchus
 
Araripesuchus
 
Elosuchus cherifiensis
 
Laganosuchus

Antaeusuchus[28]

A. taouzensis

Paired mandibles and a partial right mandible

A peirosaurid.

Araripesuchus

A. rattoides

Douira Formation

Elosuchus

E. cherifiensis

  • Gara Sbaa Formation
  • Douira Formation

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

Hamadasuchus

H. rebouli

  • Gara Sbaa Formation
  • Douira Formation?

A peirosaurid.

Kemkemia

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]

Laganosuchus

L. thaumastos

A stomatosuchid.

Lavocatchampsa[31]

Lavocatchampsa sigogneaurusselae

Anterior portion of a rostrum with mandible, with an almost complete dentition[31]

A candidodontid notosuchian.[31]

Dinosaurs

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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.

Ornithischians

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

Ankylosauria[2][32]

Indeterminate

Douira Formation

An isolated tooth.[2]

A probable ankylosaur[32]

Ornithopoda[33]

Indeterminate

Douira Formation

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

Comparable in size and shape to tracks typically attributed to Iguanodon.[33]

Sauropods

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

Rebbachisaurus

R. garasbae

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

Gara Sbaa Formation

A rebbachisaurid.

Somphospondyli[34]

Indeterminate

Anterior dorsal vertebra, partial right ischium[34]

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]

Titanosauria[35][2]

Indeterminate

  • Ifezouane Formation
  • Douira Formation

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

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

Abelisauridae[36]

Indeterminate

Isolated teeth.[36][37]
Partial right femur.[4] Partial right maxilla.[2] A cervical vertebra.[38] Left maxilla.[2]

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

 
Carcharodontosaurus saharicus
 
Deltadromeus agilis
 
Rebbachisaurus garasbae
 
Rugops primus
 
Spinosaurus aegyptiacus reconstruction

Carcharodontosaurus[1]

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

Deltadromeus

D. agilis

Gara Sbaa Formation

"Partial skeleton, isolated limb elements."[41]

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

Dromaeosauridae[36]

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.

Noasauridae[38]

Indeterminate

An anterior cervical vertebra[38]
Isolated teeth[37]

"Osteoporosia"[42] "O. gigantea"[42] A tooth and a possible neural arch from another specimen.[42] A theropod, possibly synonymous with Sauroniops.[43]

Saurischia[44]

Indeterminate

An isolated cervical vertebra.[44]

An indeterminate saurischian.

Sauroniops[45]

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]

Spinosaurus

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

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

Afrotapejara[48]

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]

 
Afrotapejara zouhri
 
Akharhynchus martilli
 
Alanqa saharica
 
Anhanguera
 
Coloborhynchus
 
Leptostomia
 
Ornithocheirus
Akharhynchus[52] A. martilli Tafilalt Ifezouane Formation A fragment of the anterior part of the premaxillae A tropeognathine anhanguerian.

Alanqa[50]

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]

Anhanguera[53]

A. cf. piscator[53]

upper Ifezouane Formation

Partial mandibular symphysis[53]

Apatorhamphus[54]

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]

Three middle cervical vertebrae.[55][50]

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]

Leptostomia[57]

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]

Nicorhynchus[56]

N. fluviferox[58][56]

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.

Ornithocheirus[53]

O. cf. simus.[53]

upper Ifezouane Formation

Premaxillae fragment[53]

Siroccopteryx[59]

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]

See also

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References

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  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. Bibcode:2020ZooK..928....1I. doi:10.3897/zookeys.928.47517. PMC 7188693. PMID 32362741.
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