Birket Qarun Formation

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]

Birket Qarun Formation
Stratigraphic range: Eocene ~41.2–33.9 Ma PreꞒ Ꞓ O S D C P T J K Pg N [1]
Type	Sedimentary
Underlies	Qasr el Sagha Formation
Overlies	Gehannam Formation
Thickness	up to 50 metres (160 ft)
Lithology
Primary	Sandstone, claystone, shale,limestone
Location
Region	Faiyum Governorate
Country	Egypt

Paleobiota

Molluscs

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

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

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

Chondrichthyes

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

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

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

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

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

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

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

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

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

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

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); the associated skull is cataloged as SMNS 11413	A Basilosauridae cetacean

Plants

Isolated Dicotyledon woods are known.^[41] Unidentified large woody roots are known.^[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
Detarioxylon[42]	aegyptiacum	Middle Member	Birket Qarun	Wood	Terrestrial Angiosperm (Fabaceae)
Nypa?[41]	fruticans?, burtini?	Middle Member	Wadi Hitan	Rhizome casts Leaf impression	Mangrove Palm (Arecaceae)	Extant representatives of the genus
Rhizophora[6][41]	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