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Birket Qarun Formation

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Birket Qarun Formation
Stratigraphic range: Eocene
~41.2–33.9 Ma [1]
TypeSedimentary
UnderliesQasr el Sagha Formation
OverliesGehannam Formation
Thicknessup to 50 metres (160 ft)
Lithology
PrimarySandstone, claystone, shale,limestone
Location
RegionFaiyum Governorate
CountryEgypt

The Birket Qarun Formation is an Eocene aged formation in Egypt. It is part of the famous Wadi al Hitan.[2] Notable fossils include the ancient whales Basilosaurus and Dorudon as well as sirenians Eotheroides and Eosiren. It also contains the teeth of various sharks and reptiles. The area was likely a mix of both marine and freshwater area with many freshwater deposits being found alongside a marine influence from the Tethys Sea.[3] Fossils here also preserve a shallow marine nursery for the whale Dorudon and it seems that this area would have also been the site of active predation from Basilosaurus as it attacked the newly born Dorudon calves.[4]

Paleobiota

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Molluscs

[edit]

Several supposed rizholiths and rizhomes have been re-interpreted as burrows made by burrowing bivalves.[5][6]

Genus Species Member Locality Material Notes Images
Amaea (Acrila)[7] niloatica Temple Member Gebel Gehannam Shells A Wentletrap
Acanthocardia (Schedocardia)[7] gehannamensis Temple Member Gebel Gehannam Shells A Carditid
Carolia[7] placunoides Temple Member Gebel Gehannam Shells A Scallop
Claibornicardia[7] pharaonum Temple Member Garet Gehannam Shells A Carditid
Cubitostrea[7] multicostata Temple Member Gebel Gehannam Shells An Oyster
Ostrea[7] reili, fraasi, roncana Temple Member Gebel Gehannam Shells An Oyster
Turritella (Torquesia)[7] pharaonica Temple Member Gebel Gehannam Shells A Turritellidae

Actinopterygians

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Genus Species Member Locality Material Notes Images
Gymnarchus[8] sp. Umm Rigl Member Birket Qarun Isolated remains A Mormyridae elephantfish
Hydrocynus[8] sp. Umm Rigl Member Birket Qarun Isolated remains An Alestid
Qarmoutus[8] hitanensis Umm Rigl Member Birket Qarun Isolated remains A sea catfish
Mochokidae[8] indeterminate Umm Rigl Member Birket Qarun Isolated remains Catfish
Perciformes[8] indeterminate Umm Rigl Member Birket Qarun Isolated remains Perch
Polypterus[8] sp. Umm Rigl Member Birket Qarun Isolated remains A Bichir

Dipnomorphs

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Genus Species Member Locality Material Notes Images
Protopterus[8] elongus Umm Rigl Member Birket Qarun Isolated tooth plates A Protopterid

Chondrichthyes

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Genus Species Member Locality Material Notes Images
Abdounia[9] sp. Temple Member Wadi Hitan Teeth A Requiem shark
Aetobatus[9] sp. Temple Member Wadi Hitan Teeth A Stingray
Alopias[9] sp. Temple Member Wadi Hitan Teeth A Thresher shark
Anoxypristis[9] mucrodens, sp. Temple Member Wadi Hitan Teeth A Sawfish
Brachycarcharias[9] cf. twiggsensis Temple Member Wadi Hitan Teeth A Sand tiger shark
Burnhamia[9] sp. Temple Member Wadi Hitan Teeth A Mobulid
Carcharias[9] aff. twiggsensis, sp. Temple Member Wadi Hitan Teeth A Sand shark
Carcharhinus[9] sp. Temple Member Wadi Hitan Teeth A Requiem shark
Chiloscyllium[9] sp. Temple Member Wadi Hitan Teeth A Carpet shark
Coupatezia[9] wousteri, attiai, sp. Temple Member Wadi Hitan Teeth A Stingray
Crassescyliorhinus[9] sp. Temple Member Wadi Hitan Teeth A Scyliorhinid
Dasyatidae[9] Indeterminate Temple Member Wadi Hitan Teeth A Whiptail stingray
Eoplinthicus[9] sp. Temple Member Wadi Hitan Teeth An Eagle ray
Galeocerdo[9] sp. Temple Member Wadi Hitan Teeth A Tiger shark
Gymnura[9] sp. Temple Member Wadi Hitan Teeth A Stingray
Hemipristis[9] sp. Temple Member Wadi Hitan Teeth A Hemigaleid
Hexanchus[9] sp. Temple Member Wadi Hitan Teeth A Sixgill shark
Himantura[9] sp. Temple Member Wadi Hitan Teeth A Stingray
Leptocharias[9] sp. Temple Member Wadi Hitan Teeth A Hemigaleid
Lophobatis[9] sp. Temple Member Wadi Hitan Teeth An Eagle ray
Macrorhizodus[9] praecursor Temple Member Wadi Hitan Teeth A Lamnid
Megascyliorhinus[9] sp. Temple Member Wadi Hitan Teeth A Scyliorhinid
Moerigaleus[10] vitreodon Temple Member Wadi Hitan Teeth A Hemigaleid
Myliobatis[9] sp. Temple Member Wadi Hitan Teeth An Eagle ray
Narcine[9] sp. Temple Member Wadi Hitan Teeth A Numbfish
Nebrius[9] sp. Temple Member Wadi Hitan Teeth A Nurse shark
Neotrygon[9] sp. Temple Member Wadi Hitan Teeth A Whiptail stingray
Negaprion[9] frequens, amekiensis, sp. Temple Member Wadi Hitan Teeth A Lemon shark
Odontorhytis[9] aff. pappenheimi Temple Member Wadi Hitan Teeth Indet. elasmobranch
Otodus[9] sokolovi Temple Member Wadi Hitan Teeth An Otodontid
Ouledia[9] sp. Temple Member Wadi Hitan Teeth A Stingray
Pastinachus[9] sp. Temple Member Wadi Hitan Teeth A Whiptail stingray
Physogaleus[9] sp. Temple Member Wadi Hitan Teeth A Requiem shark
Platyrhina[9] sp. Temple Member Wadi Hitan Teeth A Guitarfish
Platyrhinoidis[9] sp. Temple Member Wadi Hitan Teeth A Guitarfish
Propristis[9] schweinfurthi, lathami, sp. Temple Member Wadi Hitan Teeth A Sawfish
Rhinoptera[9] sp. Temple Member Wadi Hitan Teeth A Cownose ray
Rhizoprionodon[9] sp. Temple Member Wadi Hitan Teeth A Sharpnose shark
Rhynchobatus[9] sp. Temple Member Wadi Hitan Teeth A Wedgefish
Squatina[9] sp. Temple Member Wadi Hitan Teeth An Angelshark
Stegostoma[9] sp. Temple Member Wadi Hitan Teeth A Zebra shark
Scyliorhinidae[9] Indeterminate Temple Member Wadi Hitan Teeth A Scyliorhinid
Torpedo[9] sp. Temple Member Wadi Hitan Teeth A Torpedinid
Triakidae[9] Indeterminate Temple Member Wadi Hitan Teeth A Houndshark
Triakis[9] sp. Temple Member Wadi Hitan Teeth A Houndshark

Crocodylomorphs

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Genus Species Member Locality Material Notes Images
Crocodyliformes[11] indeterminate Umm Rigl Member Birket Qarun DPC 24618 and 21120 Indeterminate Crocodrile material
Gavialoidea[11] indeterminate Umm Rigl Member Birket Qarun DPC 24708 and 24343 Indeterminate Crocodrile material
Sebecosuchia[11] indeterminate Umm Rigl Member Birket Qarun DPC 20814 Indeterminate Ziphosuchian material
Tomistoma[12] kerunense Temple Member Birket-el-Qurun CGM 9350 - type (skull) A Tomistomine

Testudines

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Genus Species Member Locality Material Notes Images
Cordichelys[13] antiqua, sp. Temple Member Wadi Hitan A partial skull, mostly complete shell, and partial pelves; UMMP 97531, a mostly complete, well-preserved lower jaw A Podocnemididae sideneck turtle
Dacquemys[14] paleomorpha Temple Member Dimeh SMNS 12645 A Podocnemididae sideneck turtle

Squamates

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Genus Species Member Locality Material Notes Images
Amphisbaenia[15] indeterminate Umm Rigl Member Birket Qarun MUVP 453, an isolated presacral vertebra Indeterminate legless squamate
Booid[16] "taxon A", "taxon B" Umm Rigl Member Birket Qarun 1 left maxilla (DPC 21241), 3 anterior trunk vertebrae (DPC 25605, 25643, and 25655), 17 mid-trunk vertebrae (DPC 25603, 25606, 25613, 25618, 25619, 25628, 25635, 25642, 25644 to 25646, 25649, 25660, 25670, 25671, 25684, and 25690), 4 posterior trunk vertebrae (DPC 25623, 25673, 25685, and 25686); 7 mid-trunk vertebrae (DPC 25610, 25614, 25615, 25629, 25639, 25651, and 25652), 5 posterior trunk vertebrae (DPC 25621, 25622, 25638, 25662, and 25672) Indeterminate boa material
Caenophidia[16] indeterminate Umm Rigl Member Birket Qarun 1 "badly eroded partial vertebra" (DPC 25679) Indeterminate snake material
Gigantophis[17] garstini Umm Rigl Member Birket Qarun 1 "giant" mid-trunk vertebra (DPC 25616), 1 "large" juvenile mid-trunk vertebra (DPC 25641) A giant madtsoiid snake
Procerophis[15] sp. Umm Rigl Member Birket Qarun MUVP 454, an isolated mid- trunk vertebra Indeterminate legless squamate
Pterosphenus[17] schweinfurthi Umm Rigl Member Birket Qarun 5 anterior trunk vertebrae (DPC 21270, 21287, 24291, 25666, and 25667), 4 mid-trunk vertebrae (DPC 25607, 25630, 25668, and 25680) A Palaeophiidae sea snake
Renenutet[16] enmerwer, sp. Umm Rigl Member Birket Qarun holotype: GCM 83731, a mid-trunk vertebra; paratypes: DPC 25657, a mid-trunk vertebra, and DPC 25627, a posterior trunk vertebra; referred specimens: 7 mid-trunk vertebrae (DPC 25609, 25611, 25633, 25640, 25656, 25657, and 25687), 2 posterior trunk vertebrae (DPC 25602 and 25608); MUVP 455, mid-trunk vertebra (4.1 mm centrum length); MUVP 457, MUVP 458, MUVP 459, anterior caudal vertebrae (5.1 mm centrum length and 4.9 mm height of the neural arch and spine); MUVP 460, posterior caudal vertebra (2 mm centrum length). A Colubroidea snake
Russellophiid[16] indeterminate Umm Rigl Member Birket Qarun 1 single mid-trunk vertebra (DPC 25663) Indeterminate snake material
Tropidophiid[16] indeterminate Umm Rigl Member Birket Qarun 1 anterior trunk vertebra (DPC 25650), 3 mid-trunk vertebrae (DPC 25634, 25654, and 25664), 2 caudal vertebrae (DPC 25674 and 25692) Indeterminate dwarf boa material
Serpentes[16] indeterminate Umm Rigl Member Birket Qarun DPC 25620, 25661, 25665 (Pterosphenus or Gigantophis?), 25677, 25678, 25689, 25691, 25693, and 2694, incomplete and/or strongly eroded vertebrae Indeterminate snake material
Varanus[18] sp. Umm Rigl Member Birket Qarun Vertebrae: DPC 21355, one cervical, two trunk and two caudal; 21442, one caudal; 23257, two trunk and two caudal; 23258, four caudals; 23778, caudal A Monitor lizard

Birds

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Genus Species Member Locality Material Notes Images
Eopelecanus[19] aegyptiacus Temple Member Garet Gehannam MUVP 505, a limb element (right tibiotarsus) An early species of pelican

Afrotheres

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Genus Species Member Locality Material Notes Images
Barytherium[20][21] grave Temple Member Birket Qarun Partial mandible and isolated teeth A member of Barytheriidae
Dilambdogale[22] gheerbranti Umm Rigl Member Birket Qarun CGM 66005 - holotype; referred material: DPC 23306E, 23307A, 23007H, 23736A, 23780C, 23784D, 23983C, 24001A, 24081B, 24103A and DPC 24108A A Tenrecomorph
Dimaitherium[23] patnaiki Umm Rigl Member Birket Qarun Holotype: CGM 42206, partial left mandible A Hyracoidean
Eosiren[24] libyca, stromeri Temple Member Birket Qarun; Wadi al Hitan Multiple articulated & isolated remains A member of Dugongidae
Eotheroides[24] clavigerum, sandersi Temple Member Birket Qarun; Wadi al Hitan Multiple articulated & isolated remains A member of Dugongidae
Moeritherium[20][21][1] lyonsi Temple Member Birket Qarun Partial mandible and isolated teeth A member of Moeritheriidae

Marsupials

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Genus Species Member Locality Material Notes Images
Ghamidtherium[25] dimaiensis Umm Rigl Member Birket Qarun GM 83699, a right dentary fragment including 2-3, and alveoli for M4 and one tooth anterior to M2 A possible Didelphimorphian

Rodents

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Genus Species Member Locality Material Notes Images
Kabirmys[26] qarunensis Umm Rigl Member Birket Qarun Isolated teeth, partial mandibles, and an edentulous partial maxilla A Nementchamyidae rodent
Protophiomys[27] aegyptensis, attiai Umm Rigl Member Birket Qarun CGM 83695 - holotype; CGM 83690 ; referred material: DPC specimens A Hystricognathi rodent
Shazurus[28] minutus Umm Rigl Member Birket Qarun CGM 83701, a left Molar; referred 15 teeth A member of Anomalure

Bats

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Genus Species Member Locality Material Notes Images
Aegyptonycteris[29] knightae Umm Rigl Member Birket Qarun CGM 83740, fragment of right maxilla with M2 and M3; only known specimen A member of Aegyptonycteridae
Qarunycteris[30] moerisae Umm Rigl Member Birket Qarun CGM 83671, a tooth (right [upper] M2) A member of Rhinopomatidae
Witwatia[30] schlosseri, eremicus Umm Rigl Member Birket Qarun GM 83668, left dentary with c1–m3; CGM 83669, left M1 or M2 A member of Philisidae

Primates

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Genus Species Member Locality Material Notes Images
Afradapis[31] longicristatus Umm Rigl Member Birket Qarun CGM 83690, partial left mandible preserving P4–M3 and masseteric fossa A member of Caenopithecinae
Biretia[32] fayumensis, megalopsis Umm Rigl Member Birket Qarun CGM 83658, mandible with m2 and m3; CGM 83661, partial mandible with m2 and m3 A Parapithecidae monkey
Karanisia[33] clarki Umm Rigl Member Birket Qarun CGM 40265, left mandible preserving M1 to M3 A possible member of Lorisidae
Masradapis[34] tahai Umm Rigl Member Birket Qarun Mandibular, maxillary fragments and isolated teeth A member of Caenopithecinae
Nosmips[35] aenigmaticus Umm Rigl Member Birket Qarun CGM 66002 - holotype A primate of uncertain affinites
Saharagalago[33] misrensis Umm Rigl Member Birket Qarun CGM 40266, a left M1 A member of Galagidae

Cetaceans

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Genus Species Member Locality Material Notes Images
Ancalecetus[36] simonsi Temple Member Wadi Hitan CGM 42290, a partial skeleton A Basilosauridae cetacean
Basilosaurus[37][38] isis, cetoides All Members Dimeh SW; Birket Qarun; Garet el Naqb; Birket-el-Qurun; Garet Gehannam; Wadi Hitan Multiple articulated & isolated remains A Basilosauridae cetacean
Dorudon[38][39] atrox All Members El Kenissa; Geziret el-Qarn; Qasr Qarun; Birket-el-Qurun; Wadi Hitan Multiple articulated & isolated remains A Basilosauridae cetacean
Masracetus[40] markgrafi Temple Member Dimeh SMNS 11414, a set of vertebrae (lumbar vertebrae) A Basilosauridae cetacean

Plants

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Isolated Dicotyledon woods are know.[41] Unidentified large woody roots are know.[6]

Genus Species Member Locality Material Notes Images
Avicennia?[6][41] marina? Middle Member
  • Minqar El-Hut
  • Sandouk El-Borneta
  • Wadi Hitan
 Type-1 rhizoliths Mangrove plant (Avicenniaceae)
Extant representatives of the genus
Cymodocea[24] sp. Temple Member Qasr Qarun
  • Foliar remains
  • Type-3 rhizoliths
 A marine Angiosperm (Cymodoceaceae)
Extant representatives of the genus
Nypa?[41] sp. Middle Member Wadi Hitan
  • Rhizome casts
  • Leaf impression
 Mangrove Palm (Arecaceae)
Extant representatives of the genus
Rhizophora[6] sp. Middle Member
  • Minqar El-Hut
  • Sandouk El-Borneta
  • Wadi Hitan
 Type-2 rhizoliths Mangrove plant (Rhizophoraceae)
Extant representatives of the genus
Thalassodendron[24] sp.
  • Middle Member
  • Temple Member
  • Qasr Qarun
  • Wadi Hitan
  • Foliar remains
  • Type-3 rhizoliths
 A marine Angiosperm (Cymodoceaceae)
Extant representatives of the genus

References

[edit]
  1. ^ a b Seiffert, Erik R.; Bown, Thomas M.; Clyde, William C.; Simons, Elwyn (2008), Fleagle, John G.; Gilbert, Christopher C. (eds.), "Geology, Paleoenvironment, and Age of Birket Qarun Locality 2 (BQ-2), Fayum Depression, Egypt", Elwyn Simons: A Search for Origins, New York, NY: Springer New York, pp. 71–86, doi:10.1007/978-0-387-73896-3_8, ISBN 978-0-387-73895-6, retrieved 2024-09-16
  2. ^ "Birket Qarun Formation". PBDB.org.
  3. ^ Murray, Alison M.; Cook, Todd D.; Attia, Yousry S.; Chatrath, Prithijit; Simons, Elwyn L. (2010-05-18). "A freshwater ichthyofauna from the late Eocene Birket Qarun Formation, Fayum, Egypt". Journal of Vertebrate Paleontology. 30 (3): 665–680. doi:10.1080/02724631003758060. ISSN 0272-4634.
  4. ^ M. Fahlke, Julia (2012-11-15). "Bite marks revisited – evidence for middle-to-late Eocene Basilosaurus isis predation on Dorudon atrox (both Cetacea, Basilosauridae)". Palaeontologia Electronica. Retrieved 2024-09-16.
  5. ^ Gee, Carole T.; Sander, P. Martin; Peters, Shanan E.; El-Hennawy, Mohamed Talaat; Antar, Mohammed Sameh M.; Zalmout, Iyad S.; Gingerich, Philip D. (2019-06-01). "Fossil burrow assemblage, not mangrove roots: reinterpretation of the main whale-bearing layer in the late Eocene of Wadi Al-Hitan, Egypt". Palaeobiodiversity and Palaeoenvironments. 99 (2): 143–158. doi:10.1007/s12549-018-0337-0. ISSN 1867-1608.
  6. ^ a b c d Abdel-Fattah, Zaki A.; Gingras, Murray K. (2020-12-15). "Origin of compound biogenic sedimentary structures in Eocene strata of Wadi El-Hitan universal heritage area, Fayum, Egypt: Mangrove roots or not?". Palaeogeography, Palaeoclimatology, Palaeoecology. 560: 110048. doi:10.1016/j.palaeo.2020.110048. ISSN 0031-0182.
  7. ^ a b c d e f g Moneer, El Sayed M.; Hewaidy, Abdel Galil A.; Ayyad, Haitham M.; Samir, Ahmed; Antar, Mohammed S.; El-Balkiemy, Arafa F.; Bazeen, Youssef S. (2024). "Taxonomy, paleoecology, and paleobiogeographical significance of the middle-late Eocene molluscs from the Garet Gehannem section in the Fayoum depression, Egypt". Journal of African Earth Sciences. 214: 105231. doi:10.1016/j.jafrearsci.2024.105231. ISSN 1464-343X.
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  9. ^ 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 Underwood, Charlie J.; Ward, David J.; King, Christopher; Antar, Sameh M.; Zalmout, Iyad S.; Gingerich, Philip D. (2011). "Shark and ray faunas in the Middle and Late Eocene of the Fayum Area, Egypt". Proceedings of the Geologists' Association. 122 (1): 47–66. doi:10.1016/j.pgeola.2010.09.004. ISSN 0016-7878.
  10. ^ Underwood, Charlie J.; Ward, David J. (2011). "New hemigaleid shark from the late Eocene of Wadi Al-Hitan, Egypt". Journal of Vertebrate Paleontology. 31 (3): 707–711. doi:10.1080/02724634.2011.563763. ISSN 0272-4634.
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  12. ^ Andrews, C. W. (1905). "I.—Notes on some New Crocodilia from the Eocene of Egypt". Geological Magazine. 2 (11): 481–484. doi:10.1017/s0016756800128456. ISSN 0016-7568.
  13. ^ Cherney, Michael D.; Wilson Mantilla, Jeffrey A.; Gingerich, Philip D.; Zalmout, Iyad; Antar, Mohammed Sameh M. (2020-11-02). "New Specimens of the Late Eocene Turtle Cordichelys (Pleurodira: Podocnemididae) From Wadi Al Hitan and Qasr El-Sagha in the Fayum Province of Eqypt". Contributions from the Museum of Paleontology, University of Michigan. 33 (2): 29–64.
  14. ^ GAFFNEY, EUGENE S.; DEBLIEUX, DONALD D.; SIMONS, ELWYN L.; SÁNCHEZ-VILLAGRA, MARCELO R.; MEYLAN, PETER A. (2002). "Redescription of the Skull of Dacquemys Williams, 1954, a Podocnemidid Side-Necked Turtle from the Late Eocene of Egyp". American Museum Novitates. 3372: 1–16. doi:10.1206/0003-0082(2002)372<0001:rotsod>2.0.co;2. ISSN 0003-0082.
  15. ^ a b El-Hares, Marwa A.; Zaher, Hussam; El-Mekkawy, Desouki; El-Sayed, Sanaa; Seiffert, Erik R.; Sallam, Hesham M. (2021-07-04). "New records of legless squamates from the lowest upper Eocene deposits of the Fayum Depression, Egypt". Journal of Vertebrate Paleontology. 41 (4). doi:10.1080/02724634.2021.1992770. ISSN 0272-4634.
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  17. ^ a b Andrews, Charles William; Andrews, Charles William (1901). "Preliminary Note on some recently discovered Extinct Vertebrates from Egypt. Part II". The geological magazine, or, Monthly journal of geology. 8: 436––444. doi:10.1017/s0016756800179750.
  18. ^ HOLMES, ROBERT B.; MURRAY, ALISON M.; ATTIA, YOUSRY S.; SIMONS, ELWYN L.; CHATRATH, PRITHIJIT (2010-09-16). "Oldest known Varanus (Squamata: Varanidae) from the Upper Eocene and Lower Oligocene of Egypt: support for an African origin of the genus". Palaeontology. 53 (5): 1099–1110. doi:10.1111/j.1475-4983.2010.00994.x. ISSN 0031-0239.
  19. ^ El Adli, Joseph J.; Wilson Mantilla, Jeffrey A.; Antar, Mohammed Sameh M.; Gingerich, Philip D. (2021-01-02). "The earliest recorded fossil pelican, recovered from the late Eocene of Wadi Al-Hitan, Egypt". Journal of Vertebrate Paleontology. 41 (1). doi:10.1080/02724634.2021.1903910. ISSN 0272-4634.
  20. ^ a b Sanders, William J.; Gheerbrant, Emmanuel; Harris, John M.; Saegusa, Haruo; Delmer, Cyrille (2010-07-20), "Proboscidea", Cenozoic Mammals of Africa, University of California Press, pp. 161–251, retrieved 2025-02-06
  21. ^ a b Alexander G S C Liu; Seiffert, Erik R; Simons, Elwyn L (2008-04-14). "Stable isotope evidence for an amphibious phase in early proboscidean evolution". Proceedings of the National Academy of Sciences of the United States of America. 105 (15). doi:10.1073/pna. Archived from the original on 2024-12-02.
  22. ^ Seiffert, Erik R. (2010). "The Oldest and Youngest Records of Afrosoricid Placentals from the Fayum Depression of Northern Egypt". Acta Palaeontologica Polonica. 55 (4): 599–616. doi:10.4202/app.2010.0023. ISSN 0567-7920.
  23. ^ Barrow, Eugenie; Seiffert, Erik R.; Simons, Elwyn L. (2010-05-21). "A primitive hyracoid (Mammalia, Paenungulata) from the early Priabonian (Late Eocene) of Egypt". Journal of Systematic Palaeontology. 8 (2): 213–244. doi:10.1080/14772010903450407. ISSN 1477-2019.
  24. ^ a b c d Zalmout, Iyad S.; Gingerich, Philip D. (2012). "Late Eocene sea cows (Mammalia, Sirenia) from Wadi al Hitan in the western desert of Fayum, Egypt". University of Michigan Papers on Paleontology. 37 (1): 1–158.
  25. ^ Sánchez-Villagra, Marcelo R.; Seiffert, Erik R.; Martin, Thomas; Simons, Elwyn L.; Gunnell, Gregg F.; Attia, Yousry (2007). "Enigmatic new mammals from the late Eocene of Egypt". Paläontologische Zeitschrift. 81 (4): 406–415. doi:10.1007/bf02990252. ISSN 0031-0220.
  26. ^ SALLAM, HESHAM M.; SEIFFERT, ERIK R.; SIMONS, ELWYN L.; BRINDLEY, CHLÖE (2010). "A large-bodied anomaluroid rodent from the earliest late Eocene of Egypt: phylogenetic and biogeographic implications". Journal of Vertebrate Paleontology. 30 (5): 1579–1593. doi:10.1080/02724634.2010.501439. ISSN 0272-4634.
  27. ^ Sallam, Hesham M.; Seiffert, Erik R.; Steiper, Michael E.; Simons, Elwyn L. (2009-09-29). "Fossil and molecular evidence constrain scenarios for the early evolutionary and biogeographic history of hystricognathous rodents". Proceedings of the National Academy of Sciences. 106 (39): 16722–16727. doi:10.1073/pnas.0908702106. ISSN 0027-8424. PMC 2757825.
  28. ^ SALLAM, HESHAM M.; SEIFFERT, ERIK R.; SIMONS, ELWYN L. (2010-07-19). "A highly derived anomalurid rodent (Mammalia) from the earliest Late Eocene of Egypt". Palaeontology. 53 (4): 803–813. doi:10.1111/j.1475-4983.2010.00962.x. ISSN 0031-0239.
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