Berberosaurus(meaning "Berberlizard ", in reference to the Berbers ofMorocco) is agenusofneotheropoddinosaur,possibly aceratosaur,from theToarcian-age (Lower Jurassic) "Toundoute Continental Series" (Azilal Formation) found in the CentralHigh Atlasof Toundoute,Ouarzazate,Morocco. Thetype speciesof the genusBerberosaurusisB. liassicus,in reference to theLiasepoch.Berberosaurusmight be the oldest known ceratosaur, and is based on partialpostcranialremains. This genus represents the oldest formally identified theropod from the North of Africa, as well one of the few from the region in the Early Jurassic.[2]

Berberosaurus
Temporal range:Toarcian
~183–174MaUncertain allocation of the layers
Liferestorationand size comparison
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Averostra
Clade: Ceratosauria
Genus: Berberosaurus
Species:
B. liassicus
Binomial name
Berberosaurus liassicus
Allainet al.,2007
Synonyms

Discovery and history

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The remains ofBerberosauruswere discovered during a series of expeditions to the High Atlas beginning in the early 2000s, when in 6 years, where made to dig in the local redbeds. It isbased onan associated partial postcranial skeleton of a subadult individual cataloged in theMuséum d'Histoire Naturelle de Marrakech;bones from this skeleton includeMHNM-Pt9a neck vertebra;MHNM-Pt23,an anterior part of thesacrum;MHNM-Pt22,a metacarpal;MHNM-Pt19,afemur;MHNM-Pt21,proximal end of the lefttibia;MHNM-Pt20,bothfibulae;MHNM-Pt16,part of anotherfemur,has been assigned to the genus as well. Its remains were found alongside those ofTazoudasaurusand an indeterminate large-bodiedtheropodwithinbone bedsinmudflowdeposits. Latertectonicactivity has affected the bones.[2]Recent papers have quoted that new material of this genus was recovered on the same area, namely the axis, a postorbital, the cranium and teeth, that are currently being studied.[3]Berberosaurus is characterized by the following features: the cervical vertebra is highly pneumatic, with short cervical centra and holes in the neural arch, with low and short neural spine, unlikeElaphrosaurusandCeratosaurus.It has anteroposteriorly short centra and neural spine; the sacral series markedly arched; the central sacrum narrow transversely; the metacarpal with a very grooved proximal end; anterior femoral trochanter reaches proximally to the midpoint of the femoral head, unlikeCeratosaurus;large femoral trochanteric platform; tibia with subtriangular distal profile; presence of an oblique ridge that covers proximally the medial sulcus of the fibula.[2]

Classification

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Diagram showing the evolution of thetheropodhand, includingBerberosaurus,as hypothesized byDal Sassoand colleagues in 2018

Ronan Allain and colleagues, who describedBerberosaurus,performed aphylogenetic analysisand found their new genus to be the mostbasalknownabelisauroid,more derived thanElaphrosaurus,Ceratosaurus,andSpinostropheus,but less so thanXenotarsosaurusandabelisaurians.Berberosaurusis distinguished from other theropods by anatomical details found in itsvertebrae,metacarpals,and hindlimb bones. Its assignment as an abelisauroid pushes back the record of this group and shows that it had diversified by the Early Jurassic.[2]However, Carrano and Sampson (2008) found it to be a basal ceratosaur outsideNeoceratosauriainstead.[4]Subsequently, the analysis of Xuet al.(2009) recovered it as adilophosauridin unresolved polytomy withDilophosaurus wetherilli,"Dilophosaurus" sinensis,DracovenatorandCryolophosaurus.[5]The phylogenetic analysis performed by Ezcurra, Agnolin and Novas (2010) recoveredBerberosaurusin unresolvedpolytomywithCeratosaurusand Abelisauroidea,[6]while the 2018 description of the basal ceratosaurSaltriovenatorplacesBerberosaurusas the sister taxon to that genus.[7]Same year, a paper found Berberosaurus in different positions: considered as a basal ceratosaurian, a neoceratosaurian or a basal abelisauroid.[8]The paper which described Berberosaurus was the last one to find a monophyletic Ceratosauria that contained both Coelophysoids and Neoceratosaurs, most papers have since excluded Coelophysoidea from Ceratosauria and instead have favored the idea that Tetanurae and Ceratosauria are both part of a monophyletic group calledAverostra,a classification which was already gaining popularity at around that time.[9]

The classification of Berberosaurus based on Delcourt et al. (2018):

Ceratosauroidea

Paleoecology and paleobiology

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Tazouda CS are a unique section due to its evidenced volcanic influence. Similar environments are found in modernPisgah cinder coneof Mojave Desert. Local floodplains alternated humid and dry periods, where in the last hypersaline Chotts and gypsium dunes likely developed

Berberosaurus,like other ceratosaurians, was abipedalcarnivore.[10]It was of moderate size; its estimated femur length of 50.5 centimetres (19.9 inches)[2]is comparable to femur lengths given byGregory S. Paulfor animals likeElaphrosaurus(52.9 cm (20.8 in))[11]andDilophosaurus(55.0 cm (21.7 in)).[12]Its remains were found with those of the earlysauropodTazoudasaurus.It has been estimated to be approximately 5 m (16 ft) long and 220–300 kg (490–660 lb) in body mass.[13][14]

The "Toundoute Continental Series", unlike other members of theAzilal Formation,due to the presence of volcanic material of coeval age. The Azilal Formation recovers a Terrestrial progradation that happened in the Central High Atlas Basin towards the Toarcian, where the older Pliensbachian Carbonate Platform retreated to the east. The lithology of this unit at Tundoute is divided in 5 units from D to H, (A-C represent the units of the underlying marine dolomite, C representing a transition to a terrestrial environment).[15]These layers have been referred to a Pliensbachian-Toarcian age, based on outdated data, as the underliying Carbonates where quoted to be a continuation with less thickess of the ones found in Todhra, that reach the Late Pliensbachian, being eveidence of a latter progradation than the one seen in the N, for example atBéni-Mellal.[16]The section including the bones ofBerberosauruswas likely deposited on a channel/floodplain type fluvial system, with sand-filled channels abundant in plant roots (mostly located in fine limestone, probably from the channel margins), developed in the near E-W direction of transit. These layers also recover the presence of thick (up to 6 m) gypsium facies, which suggest the presence of a localChott,indicating a succession of humid and dry seasons.[15]Based on mesofossils, local vegetation was apparently dominated by ferns, cycads and conifers.[15]Apart fromBerberosaurus,Tazoudasaurusand a large-bodied theropod of uncertain affinities, a possibleCoelophysidaewith juvenile & adult specimens and a small sauropod have been recovered from Acforcid, E ofDemnate,as well Gravisaurian (Tazoudasaurus?[17]) remains at the E of Azilal village, Medium-Sized Sauropod remains at Mizaguène Hill (SW Azilal) and indeterminate Dinosaur remains from other locations around Azilal and Demnate.[18]

See also

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References

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  1. ^Torices, A. (2013)."Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain".Acta Palaeontologica Polonica.doi:10.4202/app.2012.0121.
  2. ^abcdeAllain, Ronan; Tykoski, Ronald; Aquesbi, Najat; Jalil, Nour-Eddine; Monbaron, Michel; Russell, Dale;Taquet, Philippe(2007)."A basal abelisauroid from the late Early Jurassic of the High Atlas Mountains, Morocco, and the radiation of ceratosaurs"(PDF).Journal of Vertebrate Paleontology.27(3): 610–624.doi:10.1671/0272-4634(2007)27[610:AADTFT]2.0.CO;2.S2CID131617581.
  3. ^Ibrahim, N.; Sereno, P. C.; Zouhri, S.; Zouhri, S. (2017)."Les dinosaures du Maroc–aperçu historique et travaux récents"(PDF).Mémoires de la Société Géologique de France.180(4): 249–284.Retrieved21 April2023.
  4. ^Carrano & Sampson, 2008. The phylogeny of Ceratosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 6, 183-236.
  5. ^Xu, X.; Clark, J.M.; Mo, J.; Choiniere, J.; Forster, C.A.; Erickson, G.M.; Hone, D.W.E.; Sullivan, C.; Eberth, D.A.; Nesbitt, S.; Zhao, Q.; Hernandez, R.; Jia, C.-K.; Han, F.-L. & Guo, Y. (2009)."A Jurassic ceratosaur from China helps clarify avian digital homologies (supplementary information)"(PDF).Nature.459(7249): 940–944.doi:10.1038/nature08124.PMID19536256.S2CID4358448.
  6. ^Ezcurra, M.D.; Agnolin, F.L.; Novas, F.E. (2010)."An abelisauroid dinosaur with a non-atrophied manus from the Late Cretaceous Pari Aike Formation of southern Patagonia"(PDF).Zootaxa.2450:1–25.doi:10.11646/zootaxa.2450.1.1.
  7. ^Dal Sasso, Cristiano; Maganuco, Simone; Cau, Andrea (2018)."The oldest ceratosaurian (Dinosauria: Theropoda), from the Lower Jurassic of Italy, sheds light on the evolution of the three-fingered hand of birds".PeerJ.6:e5976.doi:10.7717/peerj.5976.PMC6304160.PMID30588396.S2CID56894865.
  8. ^Delcourt, R. (2018)."Ceratosaur palaeobiology: new insights on evolution and ecology of the southern rulers".Scientific Reports.8(1): 9730.Bibcode:2018NatSR...8.9730D.doi:10.1038/s41598-018-28154-x.PMC6021374.PMID29950661.
  9. ^Rauhut and Foth (2020).The Evolution of Feathers From Their Origin to the Present: From Their Origin to the Present.Fascinating Life Sciences (1 ed.). Springer.doi:10.1007/978-3-030-27223-4.ISBN9783030272227.S2CID212669998.
  10. ^Tykoski, Ronald B.; Rowe, Timothy (2004). "Ceratosauria". InWeishampel, David B.;Dodson, Peter;Osmólska, Halszka(eds.).The Dinosauria(Second ed.). Berkeley: University of California Press. pp. 47–70.ISBN0-520-24209-2.
  11. ^Paul, Gregory S.(1988).Predatory Dinosaurs of the World.New York: Simon & Schuster. pp.266.ISBN0-671-61946-2.
  12. ^Paul, Gregory S. (1988).Predatory Dinosaurs of the World.268.
  13. ^Paul, Gregory S. (2016).The Princeton Field Guide to Dinosaurs.Princeton University Press. p. 81.ISBN978-1-78684-190-2.OCLC985402380.
  14. ^Molina-Pérez & Larramendi (2016).Récords y curiosidades de los dinosaurios Terópodos y otros dinosauromorfos.Barcelona, Spain: Larousse. p. 255.
  15. ^abcMontenat, C.; Monbaron, M.; Allain, R.; Aquesbi, N.; Dejax, J.; Hernandez, J.; Taquet, P. (2005)."Stratigraphie et paléoenvironnement des dépôts volcano-détritiques à dinosauriens du Jurassique inférieur de Toundoute (Province de Ouarzazate, Haut-Atlas–Maroc)"(PDF).Eclogae Geologicae Helvetiae.98(2): 261–270.doi:10.1007/s00015-005-1161-x.S2CID129577717.RetrievedJanuary 25,2022.
  16. ^Ettaki, M.; Chellaï, E. H. (2005)."Le Toarcien inférieur du Haut-Atlas de Todrha-Dadès (Maroc): sédimentologie et lithostratigraphie".C. R. Géosciences, Paris.337(1): 814–823.Bibcode:2005CRGeo.337..814E.doi:10.1016/j.crte.2005.04.007.Retrieved25 January2022.
  17. ^Allain, R. (2012).Histoire des dinosaures.Paris: Perrin. p. 112.ISBN978-2081353053.Retrieved25 January2022.
  18. ^Jenny, J.; Jenny-Deshusses, C.; Le Marrec, A.; Taquet, P. (1980)."Découverte d'ossements de Dinosauriens dans le Jurassique inférieur (Toarcien) du Haut Atlas central (Maroc) [Discovery of dinosaur bones in the Lower Jurassic (Toarcian) of the central High Atlas (Morocco)]".Comptes Rendus de l'Académie des Sciences, Série D.290(1): 839–842.Retrieved25 January2022.