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Akrabou Formation

The Akrabou Formation is a Late Cretaceous (latest Cenomanian to early/mid Turonian)-aged geological formation and Konservat-Lagerstätte in Morocco.[1] It overlies the slightly older freshwater deposits of the Kem Kem Group, which it was once thought to be apart of. It was deposited over following the Kem Kem ecosystem's submergence by the Tethys Ocean during a marine transgression from the Cenomanian-Turonian boundary event, as part of a wider deposition of carbonate platforms across the region from the event.[2][3][4]

Akrabou Formation
Stratigraphic range: latest Cenomanian to ?middle Turonian
~93.9–90 Ma
↓
TypeGeological formation
Sub-unitsAgoult, Goulmima, others
OverliesKem Kem Group
Lithology
PrimaryLimestone
Location
Coordinates30°30′N 4°48′W / 30.5°N 4.8°W / 30.5; -4.8
Approximate paleocoordinates17°48′N 4°48′W / 17.8°N 4.8°W / 17.8; -4.8
RegionEr Rachidia
Country Morocco,  Algeria
Extentcentral and eastern Morocco north and south of the Pre-African Trough
Type section
Named byEl Mostafa Ettachfini, Bernard Andreu
Year defined2004
Akrabou Formation is located in Morocco
Akrabou Formation
Akrabou Formation (Morocco)

Localities

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The Akrabou Formation has outcrops across the Anti-Atlas, but it has two especially prominent localities: Agoult and Goulmima (also known as Asfla). The Agoult locality is also commonly referred to as Gara Sbaa due to it outcropping at the hill of the same name, but this name is also used for the underlying freshwater-based deposit of the Kem Kem Group at the same locality, which may lead to nomenclatural confusion.[5][6] Both localities are considered lagerstätten due to their excellently preserved fossils, but they differ in time period, paleoenvironment, and preservation of fossils.

The Agoult locality is the older of the two and appears to have been deposited around the time of the Cenomanian-Turonian boundary event, dating it to latest Cenomanian or earliest Turonian. It is a plattenkalk that contains excellently preserved two-dimensional fossils of fish, land plants, marine and terrestrial invertebrates, and some terrestrial reptiles. The presence of a significant terrestrial component indicates it was deposited in a shallow marine environment close to shore, although the deposits record the gradual deepening of the habitat as the sea level rose.[7]

In contrast, the Goulmima locality is younger, dating to the early or mid Turonian, and appears to have been deposited in an offshore environment on the continental shelf. Fossils from this locality, which include fish and marine reptiles, are found in concretions and have a unique three-dimensional preservation akin to that of the older Santana Formation in Brazil.[7][8]

In addition, several other localities are also known, some of which cross the border into Algeria, which have a rich fauna of ammonites and nautiloids.[9]

Paleoenvironment

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In the basal Agoult horizons of the formation, a number of plant remains are known, which are similar to those found in southern Europe at the same time, suggesting floristic exchanges between northern Africa and southern Europe. Based on these, the paleoclimate of the ecosystem, at least earlier during its deposition, is suggested to have been a mildly dry subtropical climate akin to that of the modern western Canary Islands.[6][7]

Paleobiota

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Most fish are documented in a species list from Amalfitano et al (2020) & Cooper et al (2023):[10]

Vertebrates

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Cartilaginous fish

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Genus Species Locality Material Notes Image
Asflapristis A. cristadentis Goulmima Incomplete three-dimensional skeletons, isolated teeth A ptychotrygonid sawskate.[11]
Cretomanta C. sp. Goulmima Isolated teeth A chondrichthyan of uncertain affinties, possibly an aquilolamnid.[8]
Haimirichia H. amonensis Agoult Partial skeleton A haimirichiid mackerel shark.[8]
Ptychodus P. maghrebianus Goulmima Dental battery, isolated teeth A ptychodontid shark.[12]  
Ptychotrygon P. rostrispatula Goulmima Incomplete skeletons A ptychotrygonid sawskate.[8]  
?Rhinobatos ?R. sp. Goulmima Incomplete cranium A presumed guitarfish.[8]  
Squalicorax S. sp. Goulmima Teeth A crow shark.[8]  
Tingitanius T. tenuimandibulus Goulmima Three-dimensionally preserved incomplete skeleton of a young individual The oldest known platyrhinid ray.[13]

Bony fish

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Genus Species Locality Material Notes Image
Acanthomorpha indet. Agoult Complete specimen A poorly-preserved acanthomorph of uncertain affinities.[14]
Agoultichthys A. chattertoni Agoult Complete specimen A macrosemiid.[15]
Agoultpycnodus A. aldrovandii Agoult Complete specimen A pycnodont.[16]
Amioidea gen. et sp. nov. 1 Agoult Undescribed amiiforms in their own family.[5][17]
Amioidea gen. et sp. nov. 2
Anomoeodus A. sp. "A" Goulmima Isolated jaw bones. A pycnodont.  
A. sp. "B"
Armigatus A. oligodentatus Agoult Complete specimens An armigatid clupeomorph.[18]  
Belonostomus B. sp. Agoult Complete specimen An aspidorhynchid.  
Araripichthys A. corythophorus Goulmima Incomplete specimens with associated skull remains An araripichthyiform.  
cf. Chanoidei indet. Agoult Complete specimen A potential chanid.[5]
Cladocyclus C. sp. Agoult An ichthyodectid.[7]
cf. Dercetis cf D. sp. Agoult Complete specimens A dercetid aulopiform.[5]
Enchodus E. sp. Goulmima Incomplete specimens, isolated crania & teeth, An enchodontid aulopiform.  
Errachidia E. pentaspinosa Agoult Complete specimen A pharmacichthyid lamprimorph.[14]
Goulmimichthys G. arambourgi Goulmima Complete specimen. A pachyrhizodontid crossognathiform.  
Grandemarinus G. gherisiensis Goulmima 3 specimens known, one complete. A gar.[8]  
Ghrisichthys G. bardacki Goulmima Isolated cranium An ichthyodectid.[19]
Homalopagus H. multispinosus Agoult Complete specimen A pharmacichthyid lamprimorph.[14]
Ichthyodectes I. sp. Goulmima Complete specimen, isolated crania An ichthyodectid.  
Lusitanichthys L. africanus Agoult Complete specimen A clupavid otophysan.[20]
Kradimus K. asflaensis Goulmima Complete specimen. A pachyrhizodontid crossognathiform.[21]
Maghrebichthys M. nelsoni Agoult Complete specimen A pycnosteroidid beardfish.[14]
Neomesturus N. asflaensis Goulmima Isolated jaw bones. A pycnodont.[22]
Osmeroides O. rheris Goulmima Isolated crania An osmeroidid albuliform.  
Paranursallia P. cavini Goulmima Isolated jaw bones. A pycnodont.[22]  
Pleuropholis P. danielae Agoult Complete specimen The last known pleuropholid.[23]  
Polazzodus P. sp. Goulmima Isolated jaw bones. A pycnodont.[22]  
Rhynchodercetis R. sp. Agoult Complete specimens. A dercetid aulopiform.[7]  
Saurorhamphus S. sp. Agoult Complete specimens. A eurypholid aulopiform.[7]
Sorbinichthys S. africanus Agoult Complete specimen. A sorbinichthyid clupeomorph.  
Teleostei indet. Agoult Complete specimen An indeterminate teleost.[5]
Thorectichthys T. marocensis Agoult Complete specimens. A paraclupeid clupeomorph.
T. rhadinus

Reptiles

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Genus Species Locality Material Notes Image
Brachauchenius B. lucasi Goulmima Partial mandible A pliosaur, the first record of this species outside North America.[24]  
Libonectes L. morgani (=L. atlasense) Goulmima Articulated skull and post-cranial skeleton. An elasmosaurid plesiosaur, the first record of this species outside North America.[25]  
Manemergus M. angirostris Goulmima Partially complete skeleton A polycotylid plesiosaur.  
Protostegidae indet. Goulmima Carapace, skull Indeterminate protostegid turtles.[3]
Tethysaurus T. nopcsai Goulmima Hundreds of specimens of different age classes, some nearly complete[26] A tethysaurine mosasaur.  
Thililua T. longicollis Goulmima Complete skull, lower jaw, and articulated vertebrae A polycotylid plesiosaur.  
Polycotylidae indet. Goulmima Skull with mandible An indeterminate polycotylid plesiosaur.[25]
Squamata indet. Agoult Complete specimen A small indeterminate terrestrial lizard, specimen now lost.[7]

Mollusks

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Based on Kennedy et al (2008) & Meister et al (2017):[27][9]

Genus Species Locality Material Notes Image
Angulithes A. mermeti Douira, Goulmima & Tazzougert Shells A hercoglossid nautiloid.
Choffaticeras C. segne Goulmima A pseudotissotid ammonite.
C. sinaiticum
Eutrephoceras E. sp. Douira & Taouz A nautilid nautiloid.  
Fagesia F. peroni Goulmima A vasoceratid ammonite.  
F. cf. tevesthensis
Hoplitoides H. mirabilis Goulmima A hoplitid ammonite.
H. gr. wortmanni
Mammites M. nodosoides Goulmima An acanthoceratid ammonite.  
Nannovascoceras N. intermedium Goulmima A vasoceratid ammonite.
Neolobites N. vibrayeanus Agoult, Tazzougert, Taouz, Tizi Momrad An engonoceratid ammonite.
Neoptychites N. cephalotus Goulmima A vasoceratid ammonite.
N. aff. hottingeri Goulmima
Romaniceras (Yubariceras) R. (Y.) reymenti Goulmima An acanthoceratid ammonite.
Thomasites T. rollandi Goulmima A vasoceratid ammonite.
Wrightoceras W. munieri Goulmima A pseudotissotid ammonite.

Arthropods

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Partially based on Garassino, Angeli & Pasini (2014):[28]

Genus Species Locality Material Notes Image
Afroapseudes A. cretacicus Agoult A tanaidacean crustacean.[29]
Amazighopsis A. cretacea An astacidean crustacean.[30]
Corazzatocarcinus C. cf. hadjoulae A necrocarcinid crab.  
Cretagalathea C. exigua A munidid squat lobster.
Cretapenaeus C. berberus A penaeid prawn.
Galathea G. sahariana A galatheid squat lobster.  
?Glyphea G. garaasbaensis A glypheid crustacean.
Mesolimulus M. tafraoutensis Two complete specimens A horseshoe crab.[31]  
?Isoptera indet. Complete specimen A potential termite.[32]
Marocarcinus M. pasinii A marocarcinid crab.
Muelleristhes M. africanus Complete specimen A porcelain crab.[28]
Palinuridae indet. Complete specimen A spiny lobster.[33]
Phagophytichnus P. ekowskii Trace fossil Feeding traces on a leaf.[32]
Polyphaga indet. Complete specimen A polyphagan beetle.[32]
Telamonocarcinus T. cf. gambalatus A dorippid crab.
Troponoma T. constricta Trace fossil Trail made by a leafminer.[6]
Unusuropode U. castroi Complete specimen A sphaeromatid isopod, first record of this species outside Brazil.[34]

Plants

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Based primarily on Krassilov & Bacchia (2013):[6]

Genus Species Locality Material Notes Image
Abietites A. (Tritaenia) cf. linkii Agoult Needles A conifer.
Barykovia B. cf. tschuckotika Leaf A potentially fagalean angiosperm.
Cocculophyllum C. cf. fucinerve Leaf A climbing angiosperm.
Coniopteris C. cf. Dicksonia mamiyai Leaflet. A fern.  
Dryophyllum D. cf. subcretaceum Leaf A potentially fagalean angiosperm.
Frenelopsis F. cf. teixeirae Shoot A cheirolepidiacean conifer.  
Garasbahia G. flexuosa Branching shoot with leaves. A cabombacean angiosperm.[6]
Pseudotorellia P. cf. ensiformis Leaf A ginkgoalean.
Sulcatocladus S. cf. robustus Shoot A conifer.
Welwitschiophyllum W. sp. Leaf A possible welsitschialean.[35]

References

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  1. ^ Ettachfini, El Mostafa; Andreu, Bernard (2004-04-01). "Le Cénomanien et le Turonien de la Plate-forme Préafricaine du Maroc". Cretaceous Research. 25 (2): 277–302. Bibcode:2004CrRes..25..277E. doi:10.1016/j.cretres.2004.01.001. ISSN 0195-6671.
  2. ^ 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.
  3. ^ a b Cavin, L.; Tong, H.; Boudad, L.; Meister, C.; Piuz, A.; Tabouelle, J.; Aarab, M.; Amiot, R.; Buffetaut, E.; Dyke, G.; Hua, S.; Le Loeuff, J. (2010-07-01). "Vertebrate assemblages from the early Late Cretaceous of southeastern Morocco: An overview". Journal of African Earth Sciences. 57 (5): 391–412. Bibcode:2010JAfES..57..391C. doi:10.1016/j.jafrearsci.2009.12.007. ISSN 1464-343X.
  4. ^ Ibrahim, Nizar; Sereno, Paul C.; Varricchio, David J.; Martill, David M.; Dutheil, Didier B.; Unwin, David M.; Baidder, Lahssen; Larsson, Hans C.E.; Zouhri, Samir; Kaoukaya, Abdelhadi (2020-04-21). "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. ISSN 1313-2989. PMC 7188693. PMID 32362741.
  5. ^ a b c d e Murray, Allison M.; Wilson, Mark V. H.; Gibb, Stacey; Chatterton, Brian D. E. (2013). "Additions to the Late Cretaceous (Cenomanian/Turonian) actinopterygian fauna from the Agoult locality, Akrabou Formation, Morocco, and comments on the palaeoenvironment". Mesozoic Fishes 5 – Global Diversity and Evolution. Verlag Dr. Friedrich Pfeil. pp. 525–548. ISBN 978-3-89937-159-8.
  6. ^ a b c d e Krassilov, V.; Bacchia, F. (2013-03-01). "New Cenomanian florule and a leaf mine from southeastern Morocco: Palaeoecological and climatological inferences". Cretaceous Research. 40: 218–226. Bibcode:2013CrRes..40..218K. doi:10.1016/j.cretres.2012.07.005. ISSN 0195-6671.
  7. ^ a b c d e f g Martill, David M.; Ibrahim, Nizar; Brito, Paulo M.; Baider, Lahssen; Zhouri, Samir; Loveridge, Robert; Naish, Darren; Hing, Richard (2011-08-01). "A new Plattenkalk Konservat Lagerstätte in the Upper Cretaceous of Gara Sbaa, south-eastern Morocco". Cretaceous Research. 32 (4): 433–446. Bibcode:2011CrRes..32..433M. doi:10.1016/j.cretres.2011.01.005. ISSN 0195-6671.
  8. ^ a b c d e f g Cooper, Samuel L. A.; Gunn, James; Brito, Paulo M.; Zouhri, Samir; Martill, David M. (2023-11-01). "A new fully marine, short-snouted lepisosteid gar from the Upper Cretaceous (Turonian) of North Africa". Cretaceous Research. 151: 105650. Bibcode:2023CrRes.15105650C. doi:10.1016/j.cretres.2023.105650. ISSN 0195-6671.
  9. ^ a b Meister, Christian; Piuz, André; Cavin, Lionel; Boudad, Larbi; Bacchia, Flavio; Ettachfini, El M.; Benyoucef, M. (2017). "Late Cretaceous (Cenomanian-Turonian) ammonites from southern Morocco and south western Algeria". Arabian Journal of Geosciences. 10 (1). Bibcode:2017ArJG...10....1M. doi:10.1007/s12517-016-2714-1. ISSN 1866-7511.
  10. ^ Amalfitano, Jacopo; Giusberti, Luca; Fornaciari, Eliana; Carnevale, Giorgio (2020-04-03). "UPPER CENOMANIAN FISHES FROM THE BONARELLI LEVEL (OAE2) OF NORTHEASTERN ITALY". Rivista Italiana di Paleontologia e Stratigrafia. 126 (2). doi:10.13130/2039-4942/13224. ISSN 2039-4942.
  11. ^ Villalobos-Segura, Eduardo; Underwood, Charlie J.; Ward, David J.; Claeson, Kerin M. (2019-11-02). "The first three-dimensional fossils of Cretaceous sclerorhynchid sawfish: Asflapristis cristadentis gen. et sp. nov., and implications for the phylogenetic relations of the Sclerorhynchoidei (Chondrichthyes)". Journal of Systematic Palaeontology. 17 (21): 1847–1870. Bibcode:2019JSPal..17.1847V. doi:10.1080/14772019.2019.1578832. ISSN 1477-2019.
  12. ^ Amadori, Manuel; Kindlimann, René; Fornaciari, Eliana; Giusberti, Luca; Kriwet, Jürgen (2022-03-01). "A new cuspidate ptychodontid shark (Chondrichthyes; Elasmobranchii), from the Upper Cretaceous of Morocco with comments on tooth functionalities and replacement patterns". Journal of African Earth Sciences. 187: 104440. Bibcode:2022JAfES.18704440A. doi:10.1016/j.jafrearsci.2021.104440. ISSN 1464-343X. PMC 7612291. PMID 35111270.
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  15. ^ Murray, Alison M.; Wilson, Mark V. H. (2009). "A New Late Cretaceous Macrosemiid Fish (Neopterygii, Halecostomi) from Morocco, with Temporal and Geographical Range Extensions for the Family". Palaeontology. 52 (2): 429–440. Bibcode:2009Palgy..52..429M. doi:10.1111/j.1475-4983.2009.00851.x. ISSN 0031-0239.
  16. ^ Taverne, Louis; Luigi, Capasso (2021). "Osteology and phylogenetic relationships of Agoultpycnodus aldrovandii gen. and sp. nov., a new pycnodont fish genus (Pycnodontidae) from the marine Upper Cretaceous of Morocco" (PDF). Geo-Eco-Trop. 45 (3): 487–495.
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  20. ^ Cavin, Lionel (1999-11-15). "A new clupavidae (teleostei, ostariophysi) from the Cenomanian of Daoura (Morocco)". Comptes Rendus de l'Académie des Sciences - Series IIA - Earth and Planetary Science. 329 (9): 689–695. Bibcode:1999CRASE.329..689C. doi:10.1016/S1251-8050(00)87647-8. ISSN 1251-8050.
  21. ^ Veysey, Alexander J.; Brito, Paulo M.; Martill, David M. (2020-10-01). "A new crossognathiform fish (Actinopterygii, Teleostei) from the Upper Cretaceous (Turonian) of southern Morocco with hypertrophied fins". Cretaceous Research. 114: 104207. Bibcode:2020CrRes.11404207V. doi:10.1016/j.cretres.2019.104207. ISSN 0195-6671.
  22. ^ a b c Cooper, Samuel L. A.; Martill, David M. (2020-12-01). "Pycnodont fishes (Actinopterygii, Pycnodontiformes) from the Upper Cretaceous (lower Turonian) Akrabou Formation of Asfla, Morocco". Cretaceous Research. 116: 104607. Bibcode:2020CrRes.11604607C. doi:10.1016/j.cretres.2020.104607. ISSN 0195-6671. PMC 7442934. PMID 32863512.
  23. ^ Brito, Paulo M.; Vullo, Romain (2023-08-01). "A new species of Pleuropholis (Teleostei: Pleuropholidae) from the Upper Cretaceous Akrabou Formation of southeastern Morocco, with comments on the evolutionary history of the genus". Cretaceous Research. 148: 105500. Bibcode:2023CrRes.14805500B. doi:10.1016/j.cretres.2023.105500. ISSN 0195-6671.
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  26. ^ Gale, A. S.; Kennedy, W. J.; Martill, D. (2017). "Mosasauroid predation on an ammonite – Pseudaspidoceras – from the Early Turonian of south-eastern Morocco". Acta Geologica Polonica. 67 (1): 31–46. Bibcode:2017AcGeP..67...31G. doi:10.1515/agp-2017-0003. ISSN 0001-5709.
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  29. ^ Pasini, Giovanni; Vega, Francisco J.; Garassino, Alessandro; Pasini, Giovanni; Vega, Francisco J.; Garassino, Alessandro (2022). "A new genus and species of Tanaidacea (Crustacea, Apseudomorpha) from the Upper Cretaceous (Cenomanian-Turonian) of Gara Sbaa, southeastern Morocco". Boletín de la Sociedad Geológica Mexicana. 74 (2): A290622. doi:10.18268/bsgm2022v74n2a290622. ISSN 1405-3322.
  30. ^ Garassino, Alessandro; Pasini, Giovanni (2018-02-05). "Amazighopsidae, a new family of decapod macruran astacideans from the late Cretaceous (Cenomanian-Turonian) of Gara Sbaa, Southeastern Morocco". Natural History Sciences. 5 (1). doi:10.4081/nhs.2018.358. ISSN 2385-0922.
  31. ^ Lamsdell, James C.; Tashman, Jessica N.; Pasini, Giovanni; Garassino, Alessandro (2020-02-01). "A new limulid (Chelicerata, Xiphosurida) from the Late Cretaceous (Cenomanian–Turonian) of Gara Sbaa, southeast Morocco". Cretaceous Research. 106: 104230. Bibcode:2020CrRes.10604230L. doi:10.1016/j.cretres.2019.104230. ISSN 0195-6671.
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  33. ^ Pasini, Giovanni; Garassino, Alessandro (2020-08-28). "A spiny lobster (Decapoda, Achelata) from the Upper Cretaceous (Cenomanian–Turonian) of Gara Sbaa, southeastern Morocco". Palaeodiversity. 13 (1): 83. doi:10.18476/pale.v13.a8. ISSN 1867-6294.
  34. ^ Corbacho, Joan; Morrison, Scott; Alonso, Máximo (December 2018). "First Mention of Unusuropode castroi Duarte & Santos, 1962 (Crustacea: Isopoda) in the Upper Cretaceous of Gara es Sbâa Lagerstätte, South-Eastern Morocco". Earth Sciences Research Journal. 7 (6): 288–292.
  35. ^ Roberts, Emily A.; Martill, David M.; Loveridge, Robert F. (2020-02-01). "Phytogeographical implications of the probable occurrence of the gnetalean plant Welwitschiophyllum in the Late Cretaceous (Cenomanian) of Africa". Proceedings of the Geologists' Association. 131 (1): 1–7. Bibcode:2020PrGA..131....1R. doi:10.1016/j.pgeola.2019.10.002. ISSN 0016-7878.
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