Ksar Met-Lili, Anoual: Late/Upper Tithonian - Early/Lower Berriasian, Morocco
collected by M. Monbaron, D. Sigogneau-Russell 1983–1988

List of taxa
Where & when
Geology
Taphonomy & methods
Metadata & references
Taxonomic list
Echinoidea
Echinoidea indet. (Leske 1778)
unclassified
aff. Enneabatrachus sp. Evans and Milner 1993
Jones et al. 2003 18 specimens
MCM 59-66. 78-80, 82-84, 87, 93-94, 207 (ilia, premaxillae, maxillae, atlas, sacral vertebrae, presacral vertebrae, scapulae)
    = Neobatrachia indet.
Agnolin 2012
Amphibia - Temnospondyli
Anura indet. (Fischer von Waldheim 1813)
Jones et al. 2003 14 specimens
MCM 70-72, 74-76, 88-92, 97-99 (2-3 morphotypes for most elements)
Aygroua anoualensis n. gen., n. sp. Jones et al. 2003
Jones et al. 2003 24 specimens
MCM 183 - holotype (proximal part of right ilium) and referred material: MCM 57, 61, 67-69, 77, 81-82, 85-86, 95-96, 208-210(ilia, premaxilla, maxillae, atlantes, sacral vertebra, presacral vertebrae, scapulae)
Gymnophiona indet. Rafinesque 1814
Sigogneau-Russell et al. 1998 3 individuals
    = Rubricacaecilia monbaroni n. gen., n. sp. Evans and Sigogneau-Russell 2001
Evans and Sigogneau-Russell 2001
MCM 171 (holotype pseudodentary), MCM 172-173 (left pseudodentaries), MCM 56 (right pseudangular), MCM 174 (partial palatine), MCM 175 (atlas), MCM 1-2, 176-181 (postatlantal vertebrae)
Amphibia - Temnospondyli - Albanerpetontidae
Albanerpetontidae indet. Fox and Naylor 1982
Broschinksi and Sigogneau-Russell 1996
    = Anoualerpeton unicus n. gen., n. sp. Gardner et al. 2003
Gardner et al. 2003
Reptilia - Ornithischia
Ornithischia indet. Seeley 1888
Sigogneau-Russell et al. 1990
3 varieties of unknown taxonomic meaning
Reptilia - Theropoda
Theropoda indet. Marsh 1881
Sigogneau-Russell et al. 1990 11 specimens
MNHN SA mcm 167, 2004/2A, 2004/2D, 2004/2C, 2004/2E, 2004/2F, 2004/2G, 2004/3B, 2004/4A, 2004/5C, 2004/5D
Reptilia
Coelurosauria indet. Huene 1914
Knoll and Ruiz-Omeñaca 2005 3 specimens
    = Maniraptoriformes indet. Holtz and Padian 1995
Knoll and Ruiz-Omeñaca 2009
MNHN SA A0, SA 2004/2B, 2004/4C
Maniraptora indet. Gauthier 1986
Knoll and Ruiz-Omeñaca 2005 9 specimens
    = Maniraptora informal indet. 1 Gauthier 1986
Knoll and Ruiz-Omeñaca 2009
"morphotype I"; MNHN SA 2004/1, mcm 153, mcm 162, mcm 166, 2004/3E, 2004/3F, 2004/4B, 2004/4D, 2004/3C
Maniraptora informal indet. 2 Gauthier 1986
Knoll and Ruiz-Omeñaca 2009 1 specimen
"morphotype II"; MNHN SA 2004/5A
Maniraptora informal indet. 3 Gauthier 1986
Knoll and Ruiz-Omeñaca 2009 1 specimen
"morphotype III"; MNHN SA 2004/4E
Reptilia - Dromaeosauridae
Velociraptorinae indet. Barsbold 1983
Knoll and Ruiz-Omeñaca 2005 2 specimens
    = Velociraptorinae informal indet. 1 Barsbold 1983
Knoll and Ruiz-Omeñaca 2009
"morphotype I"; SA mcm 168, 2004/3A
Velociraptorinae informal indet. 2 Barsbold 1983
Knoll and Ruiz-Omeñaca 2009 1 specimen
"morphotype II"; MNHN SA 2004/5B
Velociraptorinae informal indet. 3 Barsbold 1983
Knoll and Ruiz-Omeñaca 2009 1 specimen
"morphotype III"; MNHN SA mcm 158
Reptilia
? Gnathosaurinae indet. Unwin 1992
Knoll 2000
Abundant small teeth
    = Pterosauria indet. Kaup 1834
Unwin et al. 2000
? Ornithocheiridae indet. Seeley 1870
Knoll 2000
Numerous robust teeth
    = Pterosauria indet. Kaup 1834
Unwin et al. 2000
Reptilia - Rhamphorhynchidae
? Rhamphorhynchidae indet. (Seeley 1870)
Lasseron et al. 2019
Reptilia
Ctenochasmatidae indet. (Nopcsa 1928)
Lasseron et al. 2019
"(Gnathosaurinae?)"
Reptilia - Crocodylia
Crocodylia indet. (Owen 1842)
Sigogneau-Russell et al. 1990
Reptilia - Sphenodontia
Sphenodontia indet. (Jaekel 1910)
Evans and Sigogneau-Russell 1997 2 specimens
"Sphenodontian B"
Reptilia
Tingitana anoualae n. gen., n. sp. Evans and Sigogneau-Russell 1997
Evans and Sigogneau-Russell 1997 1 specimen
Reptilia - Squamata
Squamata indet. Oppel 1811
Lasseron et al. 2019
Reptilia - Paramacellodidae
Paramacellodidae indet. Estes 1983
Broschinksi and Sigogneau-Russell 1996 1 specimen
Paramacellodus marocensis n. sp. Richter 1994
Richter 1994 3 specimens
MCM 116 (holotype), left dentary with 12 teeth; MCM 117, fragment of right dentary; MCM 118, fragment of left dentary with 3 teeth
Reptilia - Lacertilia
Lacertilia indet. Owen 1842
Broschinksi and Sigogneau-Russell 1996 1 specimen
Reptilia
Tarratosaurus anoualensis n. gen., n. sp. Broschinksi and Sigogneau-Russell 1996
Broschinksi and Sigogneau-Russell 1996 3 specimens
Reptilia - Testudines
Pleurodira indet. Cope 1865
Sigogneau-Russell et al. 1998
Reptilia - Testudines - Araripemydidae
? Araripemydidae indet. Price 1973
Lasseron et al. 2019
aff. Taquetochelys sp. Broin 1980
Sigogneau-Russell et al. 1998
Reptilia - Testudines
Cryptodira indet. Cope 1868
Sigogneau-Russell et al. 1998
Mammalia - Hahnodontidae
Hahnodontidae indet. Sigogneau-Russell 1991
Hahn and Hahn 2003 2 specimens
MNHN SA 152, 102
Denisodon moroccensis n. gen., n. sp. Hahn and Hahn 2003
Hahn and Hahn 2003 1 specimen
MNHN SA 97
Hahnodon taqueti n. gen., n. sp. Sigogneau-Russell 1991
Sigogneau-Russell 1991 1 specimen
Mammalia - Dryolestidae
Dryolestidae indet. Marsh 1879
Sigogneau-Russell et al. 1990 1 specimen
Mammalia - Dryolestida - Donodontidae
Donodon prescriptoris n. gen., n. sp. Sigogneau-Russell 1991
Sigogneau-Russell 1991 3 specimens
MNHN SA 31 (holotype right upper molar), SA 1, SA 44
Mammalia
Zatheria indet. McKenna 1975
Lasseron et al. 2019
Mammalia - Peramura - Peramuridae
? Peramuridae indet. Kretzoi 1946
Sigogneau-Russell et al. 1990 1 specimen
Afriquiamus nessovi n. gen., n. sp. Sigogneau-Russell 1999
Sigogneau-Russell 1999 1 specimen
SA 84 (holotype upper left molar)
Peramus sp. Owen 1871
Sigogneau-Russell 1999 1 specimen
SA 37 (small tooth)
Mammalia
Tribotherium africanum n. gen., n. sp. Sigogneau-Russell 1991
Sigogneau-Russell 1991 5 specimens
MNHN SA 39 (holotype partial right upper molar) and specimens referred by Sigogneau-Russell (1995): SA 72, 56 (teeth), 71, 56 (tooth fragments)
? Tribotherium africanum Sigogneau-Russell 1991
Sigogneau-Russell 1995 1 specimen
SA 74 (left lower molar)
Tribotheria indet. Butler 1978
Sigogneau-Russell 1995 2 specimens
SA 55 (left trigonid), 4 (upper left labial tooth fragment)
? Hypomylos sp. Sigogneau-Russell 1992
Sigogneau-Russell 1995 3 specimens
SA 82 (right lower molar), 83 (right lower tooth), 45 (right incomplete trigonid)
Hypomylos phelizoni n. gen., n. sp. Sigogneau-Russell 1992
Sigogneau-Russell 1992 1 specimen
MNHN SA 54 (holotype lower right molar)
Hypomylos micros n. sp. Sigogneau-Russell 1995
Sigogneau-Russell 1995 1 specimen
SA 76 (small lower right molar; holotype)
Minimus richardfoxi n. gen., n. sp. Sigogneau-Russell 1999
Sigogneau-Russell 1999 3 specimens
SA 122 (holotype left lower molar), 10 (right lower molar), 64 (left lower molar)
Mammalia - Amphilestidae
Amphilestidae indet. Osborn 1888
Sigogneau-Russell et al. 1990 2 specimens
spelled with current rank as Amphilestidae
Mammalia
Microderson laaroussii n. gen., n. sp. Sigogneau-Russell 1991
Sigogneau-Russell 1991 1 specimen
MNHN SA 40 (holotype upper right molar)
Atlasodon monbaroni n. gen., n. sp. Sigogneau-Russell 1991
Sigogneau-Russell 1991 1 specimen
MNHN SA 27 (holotype right upper molar)
Mammalia - Eutriconodonta - Gobiconodontidae
? Gobiconodontidae indet. Jenkins and Schaff 1988
Sigogneau-Russell et al. 1990 1 specimen
Mammalia - Eutriconodonta - Triconodontidae
? Triconodontidae indet. Marsh 1887
Sigogneau-Russell et al. 1990 5 specimens
Ichthyoconodon jaworowskorum n. gen., n. sp. Sigogneau-Russell 1995
Sigogneau-Russell 1995 2 specimens
MNHN SA 46 (holotype), SA 78 (right lower molars)
Mammalia - Eutriconodonta
Dyskritodon amazighi n. gen., n. sp. Sigogneau-Russell 1995
Sigogneau-Russell 1995 1 specimen
MNHN SA 92 (holotype right lower molar)
Mammalia - Thereuodontidae
Thereuodon dahmani n. sp. Sigogneau-Russell 1989
Sigogneau-Russell and Ensom 1998
Dipnoi - Ptychoceratodontidae
Ptychoceratodontidae indet. Martin 1982
Sigogneau-Russell et al. 1990
Coelacanthimorpha - Actinistia
Actinistia indet. (Cope 1870)
Sigogneau-Russell et al. 1998
Actinopteri - Osteoglossiformes
Osteoglossiformes indet. Berg 1940
Sigogneau-Russell et al. 1998
Actinopteri - Cypriniformes
Cypriniformes indet. Bleeker 1859
Sigogneau-Russell et al. 1998
Actinopteri
Ginglymodi indet. (Cope 1871)
Lasseron et al. 2019
Actinopteri - Lepisosteiformes
cf. Lepidotes sp. Agassiz 1832
Sigogneau-Russell et al. 1990
Actinopteri - Semionotiformes - Semionotidae
Semionotidae indet. Woodward 1890
Evans and Sigogneau-Russell 1997
Actinopteri - Pycnodontiformes - Pycnodontidae
Pycnodontidae indet. (Agassiz 1833)
Evans and Sigogneau-Russell 1997
Chondrichthyes - Hybodontiformes - Lonchidiidae
Lonchidion (Lissodus) marocensis (Duffin and Sigogneau-Russell 1993)
Lasseron et al. 2019
Chondrichthyes - Hybodontiformes - Hybodontidae
Hybodus sp. Agassiz 1834
Evans and Sigogneau-Russell 1997
Egertonodus basanus (Egerton 1845)
Lasseron et al. 2019
Chondrichthyes - Hybodontiformes - Polyacrodontidae
Polyacrodus sp. Jaekel 1889
Evans and Sigogneau-Russell 1997
unclassified
Brachiopoda indet. Cuvier 1805
Bivalvia
Lamellibranchia indet.
replaced by Autobranchia
Anthozoa
Anthozoa indet. Ehrenberg 1834
"corals"
unclassified
Gymnospermophyta indet. Cleal 1994
Sigogneau-Russell et al. 1998 2 specimens
"fossil wood"
see common names

Geography
Country:Morocco State/province:Oriental County:Figuig
Coordinates: 32.7° North, 3.2° West (view map)
Paleocoordinates:25.6° North, 1.6° East
Basis of coordinate:based on nearby landmark
Geographic resolution:small collection
Time
Key time interval:Late/Upper Tithonian - Early/Lower Berriasian
Age range of interval:146.30000 - 142.70000 m.y. ago
Stratigraphy
Formation:Ksar Metlili
Stratigraphic resolution:group of beds
Stratigraphy comments: originally labeled "Couches Rouges"; dated from nannoliths and charophyte

Age is debated, usually noted as ?Berrisian or Late Tithonian–Early Berrisian based on Microvertebrate analysis (Lasseron et al. 2019)
Lithology and environment
Primary lithology:lenticular,red calcareous sandstone
Includes fossils?Y
Lithology description: "calcareous lens...about 20 cm thick"
Environment:deltaic indet.
Taphonomy
Modes of preservation:body
Degree of concentration:concentrated
Size of fossils:macrofossils,mesofossils
Preservation of anatomical detail:variable
Articulated whole bodies:none
Associated major elements:none
Disassociated major elements:many
Disassociated minor elements:many
Fragmentation:frequent
Spatial resolution:parautochthonous
Collection methods and comments
Collection methods:bulk,chemical,sieve,field collection
Rock censused:700 kg
Reason for describing collection:general faunal/floral analysis
Museum repositories:MNHN
Collectors:M. Monbaron, D. Sigogneau-Russell Collection dates:1983–1988
Metadata
Also known as:Ksar Metlili, KM 1983
Database number:45726
Authorizer:M. Carrano, J. Alroy, R. Benson, T. Cleary, P. Mannion, R. Butler Enterer:M. Carrano, R. Butler, P. Mannion, R. Benson, T. Cleary, J. Alroy
Modifier:G. Varnham Research group:vertebrate
Collections that are a subset of this one:204918, 204924, 204936, 204939, 204979, 204990
Created:2004-11-29 08:20:17 Last modified:2022-02-03 04:46:30
Access level:the public Released:2004-11-29 08:20:17
Creative Commons license:CC BY
Reference information

Primary reference:
7910.5% 18780D. Sigogneau-Russell, M. Monbaron, and D. E. Russell. 1988. Decouverte de mammiferes dans le Mesozoique moyen d'Afrique [Discovery of mammals in the middle Mesozoic of Africa]. Comptes Rendus de l'Academie des Sciences, Serie 2 307(8):1045-1050 [J. Alroy/J. Wertheim/J. Alroy]

Secondary references:

73340 F. Agnolin. 2012. A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position. Studia Geologica Salmanticensia 48(2):129-178 [P. Mannion/P. Mannion]
12097ETE A. Broschinksi and D. Sigogneau-Russell. 1996. Remarkable lizard remains from the Lower Cretaceous of Anoual (Morocco). Annales de Paléontologie (Vert.–Invert.) 82(3):147-175 [M. Carrano/M. Carrano/M. Carrano]
19461ETE F. de Broin, S. Barta-Calmus, L. Beauvais, G. Camoin, J. Dejax, M. Gayet, J.-G. Michard, T. Olivaux, J. Roman, D. Sigogneau-Russell, P. Taquet and S. Wenz. 1991. Paléobiogéographie de la Téthys: apports de la paléontologie à la localisation des rivages, des aires émergées et des plates-formes au Jurassique et au Crétacé [Paleobiogeography of the Tethys: contribution of paleontology to the location of shores, emerged lands, and marine platforms in the Jurassic and Cretaceous]. Bulletin de la Société Géologique de France, 8e série 162(1):13-26 [M. Carrano/M. Carrano]
12095ETE S. E. Evans and D. Sigogneau-Russell. 1997. New sphenodontians (Diapsida: Lepidosauria: Rhynchocephalia) from the Early Cretaceous of North Africa. Journal of Vertebrate Paleontology 17(1):45-51 [M. Carrano/M. Carrano/R. Benson]
38086 S. E. Evans and D. Sigogneau-Russell. 2001. A stem-group caecillian (Lissamphibia: Gymnophiona) from the Lower Cretaceous of North Africa. Palaeontology 44(2):259-273 [R. Benson/R. Benson]
52039 J. D. Gardner and M. Böhme. 2008. Review of the Albanerpetontidae (Lissamphibia), with comments on the paleoecological preferences of European Tertiary albanerpetontids. In J. T. Sankey & S. Baszio (ed.), Vertebrate Microfossil Assemblages: Their Role in Paleoecology and Paleobiogeography 178-218 [M. Carrano/M. Carrano]
30986 J. D. Gardner, S. E. Evans, and D. Sigogneau-Russell. 2003. New albanerpetontid amphibians from the Early Cretaceous of Morocco and Middle Jurassic of England. Acta Palaeontologica Polonica 48(2):301-319 [J. Alroy/J. Alroy]
57820 H. Haddoumi, R. Allain, S. Meslouh, G. Metais, M. Monbaron, D. Pons, J.-C. Rage, R. Vullo, S. Zouhri and E. Gheerbrant. 2016. Guelb el Ahmar (Bathonian, Anoual Syncline, eastern Morocco): first continental flora and fauna including mammals from the Middle Jurassic of Africa. Gondwana Research 29:290-319 [P. Barrett/T. Cleary]
34636ETE G. Hahn and R. Hahn. 2003. New multituberculate teeth from the Early Cretaceous of Morocco. Acta Palaeontologica Polonica 48(3):349-356 [M. Carrano/M. Carrano]
37819 M. E. H. Jones, S. E. Evans, and D. Sigogneau-Russell. 2003. Early Cretaceous frogs from Morocco. Annals of Carnegie Museum 72(2):65-97 [P. Mannion/P. Mannion]
12094ETE F. Knoll. 2000. Pterosaurs from the Lower Cretaceous (?Berriasian) of Anoual, Morocco. Annales de Paléontologie 86(3):157-164 [M. Carrano/M. Carrano/M. Carrano]
14819ETE F. Knoll and J. I. Ruiz-Omeñaca. 2005. Theropod teeth from the Berriasian of Anoual (Morocco). Journal of Vertebrate Paleontology 25(3, suppl.):78A [M. Carrano/M. Carrano]
32534ETE F. Knoll and J. I. Ruiz-Omeñaca. 2009. Theropod teeth from the basalmost Cretaceous of Anoual (Morocco) and their palaeobiogeographical significance. Geological Magazine 146(4):602-616 [M. Carrano/M. Carrano]
70302 M. Lasseron, R. Allain, E. Gheerbrant, H. Haddoumi, N.-E. Jalil, G. Métais, J.-C. Rage, R. Vullo, and S. Zouhri. 2019. New data on the microvertebrate fauna from the Upper Jurassic or lowest Cretaceous of Ksar Metlili (Anoual Syncline, eastern Morocco). Geological Magazine 1-26 [T. Cleary/T. Cleary]
70734 J. Le Loeuff, E. Läng, L. Cavin and E. Buffetaut. 2012. Between Tendaguru and Bahariya: on the age of the Early Cretaceous dinosaur sites from the Continental Intercalaire and other African formations. Journal of Stratigraphy 36(2):486-502 [M. Carrano/M. Carrano]
58341 A. Richter. 1994. Lacertilia aus der Unteren Kreide von Una und Galve (Spanien) und Anoual (Marokko). Berliner Geowissenschaftliche Abhandlungen Reihe E 14:1-147 [P. Barrett/T. Cleary]
39256 D. Sigogneau-Russell. 1991. Nouveaux Mammiferes theriens du Cretace inferieur du Maroc. Comptes-Rendus de l'Academis des Sciences du Paris, series II 313:279-285 [R. Benson/R. Benson/R. Benson]
12096ETE D. Sigogneau-Russell. 1991. First evidence of Multituberculata (Mammalia) in the Mesozoic of Africa. Neues Jahrbuch für Geologie und Paläontologie Monatshefte 1991(H2):119-125 [M. Carrano/M. Carrano/M. Carrano]
39255 D. Sigogneau-Russell. 1991. Decouverte du premier mammifere tribosphenique du Mesozoique africain. Comptes-rendus de l'Academie des Sciences de Paris, serie II 313:1635-1640 [R. Benson/R. Benson]
38328 D. Sigogneau-Russell. 1992. Hypomylos phelizoni nov. gen. nov. sp., une étape précoce de l'évolution de la molaire tribosphénique (Crétacé basal du Maroc). Geobios 25(2):389-393 [R. Benson/R. Benson]
39267 D. Sigogneau-Russell. 1995. Further data and reflexions on the tribosphenid mammals (Tribotheria) from the Early Cretaceous of Morocco. Bull Mus natl Hist nat, Paris, 4 ser 16(2-4):291-312 [R. Benson/R. Benson]
38855 D. Sigogneau-Russell. 1995. Two possibly aquatic triconodont mammals from the Early Cretaceous of Morocco. Acta Palaeontologica Polonica 40(2):149-162 [R. Benson/R. Benson/R. Benson]
39266 D. Sigogneau-Russell. 1999. Reevaluation des Peramura (Mammalia, Cladotheria) sur la base de nouveaux specimens de Cretace inferieur d'Angleterre et du Maroc. Geodiversitas 21(1):93-127 [R. Benson/R. Benson/R. Benson]
9437 D. Sigogneau-Russell and P. Ensom. 1998. Thereuodon (Theria, Symmetrodonta) from the Lower Cretaceous of North Africa and Europe, and a brief review of symmetrodonts. Cretaceous Research 19:445-470 [J. Alroy/E. Leckey/R. Benson]
14076ETE D. Sigogneau-Russell, S. E. Evans, J. F. Levine and D. A. Russell. 1998. The Early Cretaceous microvertebrate locality of Anoual, Morocco: a glimpse at the small vertebrate assemblages of Africa. In S. G. Lucas, J. I. Kirkland, and J. W. Estep (eds.), Lower and Middle Cretaceous Terrestrial Ecosystems, New Mexico Museum of Natural History and Science Bulletin 14:177-181 [M. Carrano/M. Carrano/M. Carrano]
12098ETE D. Sigogneau-Russell, M. Monbaron, and E. Kaenel. 1990. Nouvelles données sur le gisement à mammifères Mésozoïques du Haut-Atlas Marocain. Géobios 23(4):461-483 [M. Carrano/M. Carrano/J. Alroy]
70903 P. Taquet. 2010. The dinosaurs of Maghreb: the history of their discovery. Historical Biology 22(1–3):88-99 [M. Carrano/M. Carrano]
31079 D. M. Unwin, J. Lü, and N. N. Bakhurina. 2000. On the systematic and stratigraphic significance of pterosaurs from the Lower Cretaceous Yixian Formation (Jehol Group) of Liaoning, China. Mitteilungen aus dem Museum für Naturkunde in Berlin, Geowissenschaftlichen Reihe 3:181-206 [R. Butler/R. Butler/R. Butler]