Ksar Metlili Formation

The Ksar Metlili Formation is a geological formation in eastern High Atlas of Morocco, it is late Tithonian to Berriasian in age. It is approximately 80 metres (260 ft) thick and primarily consists of mudstone and sandstone, with thin calcareous beds.^[1] One of these calcareous beds near the middle of the sequence is an important microvertebrate locality.^[2] Subsequent to the original site, several other localities have been sampled. The depositional environment is thought to be near shore deltaic.^[3][4]

Ksar Metlili Formation
Stratigraphic range: Late Tithonian to Early Berriasian PreꞒ Ꞓ O S D C P T J K Pg N
Type	Geological formation
Unit of	Moroccan Red Beds
Underlies	Dekkar Group
Overlies	Anoual Formation
Thickness	~80 m (260 ft)
Lithology
Primary	Mudstone, sandstone
Other	Limestone, marl, calcarenite
Location
Coordinates	32.7°N 3.1°W / 32.7; -3.1
Approximate paleocoordinates	25.1°N 1.9°E / 25.1; 1.9
Region	Figuig Province
Country	Morocco
Extent	High Atlas
Type section
Named for	Ksar Met Lili Fort
Ksar Metlili Formation (Morocco)

Fossil content

Fish

Genus	Species	Location	Material	Notes	Images
Cypriniformes[3]	Indeterminate	Ksar Metlili	Teeth	Bony Fish
Egertonodus[3][5]	E. basanus	Ksar Metlili	16 isolated Teeth KM-A2-26, teeth KM-D1-22, teeth KM-D2-25, tooth	Hybodontid
Ginglymodi[3]	Indeterminate	Ksar Metlili	KM-A1-41, fin rays KM-A1-42, teeth; 46, scales; 47, scale; 49, scales; 50, scales KM-B'-58 and 60, 66, all ichthyoliths KM-B'-59, 61, 64-65, 72, 84, all scales KM-A2-22, mandibular fragments KM-A2-23, 27, both scales; 32. teeth KM-C-6, mandibular fragments KM-C-7, scales; 9, teeth KM-D1-21, teeth KM-D2-30, teeth	Bony Fish
Hybodus[3][6]	sp.	Ksar Metlili	Teeth	Hybodontid	reconstruction of Hybodus
Lepidotes[3]	sp.	Ksar Metlili	Teeth	Lepidotidae	reconstruction of Lepidotes
Lonchidion[5]	L. (Lissodus) marocensis	Ksar Metlili	MCM108, isolated tooth	Lonchidiid
Mawsoniidae[3]	Indeterminate	Ksar Metlili	KM-A2-11, left postparietal shield	Coelacanth
Osteoglossiformes[3]	Indeterminate	Ksar Metlili	KM-B'-89, teeth	Bony Fish
Polyacrodus[3][6]	sp.	Ksar Metlili	Teeth	Polyacrodontid
Ptychoceratodontidae[3][7]	Indeterminate	Ksar Metlili	KM-B'-56, teeth KM-A2-24, teeth KM-D1-20, teeth KM-D2-22, teeth KM-D2-22, tooth	Lungfish
Pycnodontidae[3]	Indeterminate	Ksar Metlili	KM-A1-58, teeth KM-A2-25, tooth KM-D1-23, teeth	Pycnodontiform
Semionotidae[3]	Indeterminate	Ksar Metlili	Teeth	Bony Fish
?Siluriformes/?Polypteriformes[3]	Indeterminate	Ksar Metlili	KM-A1-40, fin spine KM-B'-57, fin spine KM-D1-19, fin spines	Bony Fish

Amphibians

Genus	Species	Location	Material	Notes	Images
Alytidae[3]	Indeterminate	Ksar Metlili	KM-B'-18, dorsal vertebrae	Toad
Anoualerpeton[3][8]	A. unicus	Ksar Metlili	Right premaxilla (Holotype) KM-B'-12-13, left dentary KM-A2-7, fused frontals KM-A2-31, dentary KM-D2-28, fragmentary dentary	Albanerpetontid	Holotype maxilla
Anura[3][9]	Indeterminate	Ksar Metlili	MCM 70-72, 74-76, 88-92, 97-99 KM-B'-15, right ilium; 16, scapula; 17, dorsal vertebrae; 27, dorsal vertebrae of a hatchling or neotenic; 86, long bone fragments KM-A2-2, radioulna KM-A2-3, coracoid; 4, femur; 5, scapula; 29, long bones KM-D1-25, long bones	Frogs
Aygroua[9]	A. anoualensis	Ksar Metlili	MCM 183 - holotype (proximal part of right ilium) MCM 57, 61, 67-69, 77, 81-82, 85-86, 95-96, 208-210(ilia, premaxilla, maxillae, atlantes, sacral vertebra, presacral vertebrae, scapulae)	Frog
Enneabatrachus[9]	aff.sp.	Ksar Metlili	ACMC60, left illium	Discoglossid Frog
Rubricacaecilia[10]	R. monbaroni	Ksar Metlili	Nearly complete right pseudodentary, partial left pseudodentaries, partial right pseudangular, partial palatine, atlas, postcranial vertebrae	Stem-caecilian	Rubricacaecilia holotype

Lepidosaurs

Genus	Species	Location	Material	Notes	Images
Lacertilia[11]	Indeterminate	Ksar Metlili	Teeth	Lizard
Paramacellodus[12]	P. marocensis	Ksar Metlili	MCM 116 (holotype), left dentary with 12 teeth MCM 117, fragment of right dentary MCM 118, fragment of left dentary with 3 teeth	Paramacellodidae Lizard
Paramacellodidae[11]	Indeterminate	Ksar Metlili	Teeth	Lizard
Scincomorpha[3]	Indeterminate	Ksar Metlili	KM-A1-7, left dentary (fragmentary) KM-B'-19, right dentary	Lizard
Squamata[3][13]	Indeterminate	Ksar Metlili	KM-A1-5, dorsal vertebra KM-A1-6, humerus, 8, dorsal vertebra KM-B'-28, dorsal vertebrae; 29, jaw fragment; 90, 92, single teeth KM-A2-6, right dentary KM-A2-28, long bones KM-D1-5, dorsal vertebra KM-D1-26, vertebrae KM-D2-27, "diverse elements"	Lizard?
Sphenodontia[3][6]	Indeterminate	Ksar Metlili	"Sphenodontian B", teeth; KM-B'-14, dorsal vertebra	Distinct from Tingitana
Tarratosaurus[3][11]	T. anoualensis	Ksar Metlili	MNHN MCM 140; KM-B'-20, right dentary	Lizard
Tingitana[6]	T. anoualae	Ksar Metlili	MCM 120, a mandible	Sphenodontid

Turtles

Genus	Species	Location	Material	Notes	Images
Araripemydidae[3]	Indeterminate	Ksar Metlili	Isolated Plastrons	Pelomedusoide Turtle
Cryptodira[13]	Indeterminate	Ksar Metlili	Isolated Plastrons	Turtle
Pleurodira[3][13]	Indeterminate	Ksar Metlili	Isolated Plastrons	Turtle
Taquetochelys[13]	sp.	Ksar Metlili	Isolated Plastrons	An Araripemydid turtle
Testudinata[3][4]	Indeterminate	Ksar Metlili	KM-A1-18, osteoderms KM-A1-20 to 24, osteoderms KM-B'-39, 41, osteoderms KM-A2-14 osteoderms KM-D1-10, osteoderms KM-D2-13, osteoderms Mo-GR.23-002, bone fragment	Turtle

Choristoderes

Genus	Species	Location	Material	Notes	Images
Choristodera[3]	Indeterminate	Ksar Metlili	KM-A1-19, dorsal vertebra KM-D1-18, vertebrae KM-D2-23 and -26 KM-B'-49, dorsal vertebrae	Similar to Cteniogenys

Crocodrylomorphs

Genus	Species	Location	Material	Notes	Images
Atoposauridae[3]	Indeterminate	Ksar Metlili	KM-B'-47, 48, A1-28, 29, 34, all teeth KM-A2-18, teeth KM-C-2, tooth KM-D1-17, teeth KM-D2-14, -15 and -18, teeth; -19, tooth	Crocodrylomorph
Goniopholididae/Pholidosauridae[4]	Indeterminate	Ksar Metlili	Mo-GR.23-001, Postcranial elements & osteoderms Mo-GR.23-001, complete skull	Crocodrylomorph
Crocodylomorpha[3]	Indeterminate	Ksar Metlili	KM-B'-40, KM-A1-25, KM-A2-13, osteoderms KM-B'-45, mandibular fragment; 46, 52, 91, A1-26, 27, 30 to 33, 35 KM-A2-15 to 17, 20 all teeth KM-D1-7 and 8, both single teeth KM-D1-11, osteoderms; 12, dorsal vertebra; 13, scapula/coracoid; 14-16, all teeth KM-D2-12, osteoderms KM-D2-16, teeth; -20, tooth	Crocodrylomorph
Teleosauridae[3]	Indeterminate	Ksar Metlili	KM-B'-51, teeth KM-B'-53, tooth KM-A1-53, tooth KM-A2-30, tooth KM-C-5, teeth KM-C-8, tooth KM-D2-29, teeth	Marine Crocodrylomorph
Theriosuchus[3]	Cf.T. sp.	Ksar Metlili	KM-B'-42, premaxillary	An Atoposaurid

Pterosaurs

Genus	Species	Location	Material	Notes	Images
Ctenochasmatidae[3]	Indeterminate	Ksar Metlili	Teeth	Pterosaur
Pterosauria[14]	Indeterminate	Ksar Metlili	KM-A1-38 to 39 KM-B'-54-55 KM-A2-21, teeth KM-D2-17 and -21, teeth Numerous robust teeth Abundant small teeth	Pterosaur
Rhamphorhynchidae[3]	Indeterminate	Ksar Metlili	Teeth	Pterosaur

Dinosaurs

Genus	Species	Location	Material	Notes	Images
Dromeosauridae[3][15]	Indeterminate	Ksar Metlili	KM-A1-10, tooth KM-B'-26, teeth KM-D1-6, tooth KM-D2-11, teeth	Coelurosaur
Maniraptoriformes[3][15]	Indeterminate	Ksar Metlili	"morphotype II" MNHN SA 2004/5A "morphotype III"; MNHN SA 2004/4E KM-B'-22, tooth KM-A2-9, teeth KM-D2-8, teeth	Coelurosaur
Ornithischia[7]	Indeterminate	Ksar Metlili	3 teeth varieties of unknown taxonomic meaning KM-A2-8, tooth	Ornithischian
Theropoda[3][15][7]	Indeterminate	Ksar Metlili	KM-A1-9, claw (pedal?) KM-A1-11, claws; 12 to 13, both teeth; 55, claw (pedal?), 56, claw (ungual?) KM-B'-21, claw; KM-B'-23-25, teeth; 94, claw (ungual?) MNHN SA mcm 167, 2004/2A, 2004/2D, 2004/2C, 2004/2E, 2004/2F, 2004/2G, 2004/3B, 2004/4A, 2004/5C, 2004/5D KM-A2-10, teeth KM-C-1, claw KM-D1-9, teeth KM-D2-9, claw KM-D2-10, teeth	Theropod
Velociraptorinae[15]	Indeterminate	Ksar Metlili	"morphotype II"; MNHN SA 2004/5B "morphotype III"; MNHN SA mcm 158	Coelurosaur

Mammals

Genus	Species	Location	Material	Notes	Images
Afriquiamus[16]	A. nessovi	Ksar Metlili	SA 84 (holotype upper left molar)	Peramurid
Atlasodon[17]	A. monbaroni	Ksar Metlili	MNHN SA 27 (holotype right upper molar)	Theriiform
Amazighodon[18]	A. orbis	Ksar Metlili	MCM 618, left lower molar KM-B’-2, right lower molar	Donodontid
Amphilestidae[7]	Indeterminate	Ksar Metlili	Teeth	Amphilestid
Anoualestes[18]	A. incidens	Ksar Metlili	KM-B’-97, left lower molar MCM 620, left lower molar MCM 623, right lower molar KM-B’-1, right lower molar KM-A1-1, left lower molar	Donodontid
Cynodontia?[19]	Indeterminate	Ksar Metlili	KM-B’-44, KM-B’-43, KM-B’-98, teeth crowns KM-?-1, almost complete ?lower tooth	Two morphotypes, non tritylodontid
Denisodon[20]	D. moroccensis	Ksar Metlili	Isolated m2 MNHN SA 97	Hahnodontidae Haramiyidan
Donodon[17][18]	D. prescriptoris, D. minor, sp.	Ksar Metlili	MNHN SA 31 (holotype right upper molar) SA 1, SA 44; MCM 595, right upper molar KM-A1-2, left upper molar KM-B'-2, left lower molar; 11, right upper molar; 95, right upper molar KM-D2-5, left upper molar	Donodontid
Dryolestidae[13]	Indeterminate	Ksar Metlili	KM-D2-4, left lower molar	Dryolestid
Dryolestoidea[3]	Indeterminate	Ksar Metlili	KM-A1-1, left lower molar KM-B'-1, left lower molar; 37, teeth; 96-97, single left lower molars	Dryolestoid
Dyskritodon[21]	D. amazighi	Ksar Metlili	MNHN SA 92 (holotype right lower molar)	Eutriconodontan	Dyskritodon holotype
Eutriconodonta[3]	Indeterminate	Ksar Metlili	KM-A1-3, tooth KM-B'-3, 6, single lower molars; 8, 30, single teeth KM-A2-1, tooth KM-D2-1, tooth	Eutriconodontan
Gobiconodon[22]	G. palaios	Ksar Metlili	SA 107, a right upper molar SA Ill, a right ?upper molar (M2?) SA 119, a right upper molar SA 130, a left upper molar (M2?) SA 138, a right upper molar SA 141, a left upper molar SA 146, anterior third of a left upper molar SA 153, anterior two-thirds of a left upper molar SA 80, a worn right upper molariform SA 93, a partial left ?upper premolar SA 129, a partial right ?upper premolar	Eutriconodontan	Gobiconodon
Hahnodon[20]	H. taqueti	Ksar Metlili	left m2	Hahnodontidae Haramiyidan
Hahnodontidae[20]	Indeterminate	Ksar Metlili	Posterior upper premolar; Lower incisor	Hahnodontidae Haramiyidan
Hypomylos[23][24]	H. phelizoni, H. micros; sp.	Ksar Metlili	SA 82 (right lower molar), 83 (right lower tooth), 45 (right incomplete trigonid) SA 76 (small lower right molar; holotype) MNHN SA 54, lower right molar KM-B'-4, left lower molar; 9-10, single right lower molars	Tribosphenidan
Ichthyoconodon[21]	I. jaworowskorum	Ksar Metlili	MNHN SA 46 (holotype) SA 78 (right lower molars)	Volaticothere	Ichthyoconodon holotype
Kryptotherium[22]	K. polysphenos	Ksar Metlili	SA 22, a right lower molar Sa 69, a right lower molar SA 70, a right lower molar SA 101, a left lower molar SA 34, anterior part of a left lower ?premolar	Eutriconodonta
Mammalia[3]	Indeterminate	Ksar Metlili	KM-A1-16, incisor tooth KM-A1-17, left dentary KM-A1-36, mandibular fragments, -37, phalanx KM-B'-31-32, both single incisor teeth; 33-34, left dentary; 35, dentary; 38, right dentary; 50, mandibular fragment KM-A2-19, mandible KM-A2-12, left dentary KM-D1-1, tooth KM-D1-2, canine; 3, incisor teeth KM-D2-2, right dentary KM-D2-3, left dentary KM-D2-7, dentaries (fragments)	Mammal
Microderson[17]	M. laaroussii	Ksar Metlili	MNHN SA 40 (holotype upper right molar)	Mammal
Minimus[16]	M. richardfoxi	Ksar Metlili	SA 122 (holotype left lower molar), 10 (right lower molar), 64 (left lower molar)	Zatherian
Peramuridae[7]	Indeterminate	Ksar Metlili	Teeth	Peramurid
Peramus[16]	sp.	Ksar Metlili	SA 37 (small tooth)	Peramurid
Stylodens[18]	S. amerrukensis	Ksar Metlili	MCM 578, right upper molar	Donodontid
Symmetrodonta[3]	Indeterminate	Ksar Metlili	KM-B'-36, teeth;	Mammal
Thereuodon[25]	T. dahmani	Ksar Metlili	Teeth	Thereuodontid Symmetrodont
Tribotheria[23][26]	Indeterminate	Ksar Metlili	SA 55 (left trigonid), 4 (upper left labial tooth fragment)	Tribosphenidan
Tribotherium[23][26]	T. africanum	Ksar Metlili	MNHN SA 39 (holotype partial right upper molar) SA 72, 56 (teeth), 71, 56 (tooth fragments) KM-B'-5, right upper molar	Tribosphenidan
Triconodontidae[7]	Indeterminate	Ksar Metlili	Teeth	Triconodontid
Zatheria[3]	Indeterminate	Ksar Metlili	Teeth	Cladotherian

Plants

Genus	Species	Location	Material	Notes	Images
Coniferophyta[4]	Indeterminate	Ksar Metlili	Branched Shoots	Conifer
Phlebopteris/Piazopteris[4]	Indeterminate	Ksar Metlili	Fronds	A Matoniaceae fern	Phlebopteris
Ruffordia?[4]	R?. sp.	Ksar Metlili	Fronds	A Schizaeaceae fern