Sannine Formation

Sannine Formation
Stratigraphic range: Late Cenomanian ~95–93.5 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Type	Geological formation
Underlies	Maameltain Formation
Overlies	Hammana Formation
Thickness	Coastal: >2,000 m (6,600 ft) Mountains: 500–700 m (1,600–2,300 ft)
Lithology
Primary	Chalk, limestone
Other	Marl
Location
Coordinates	Sannine Formation (Lebanon)

The Sannine Formation, also called the Sannine Limestone, is a Cretaceous geologic formation in Lebanon. It is a Konservat-Lagerstätte that contains a high diversity of well-preserved fish, reptiles, and invertebrates within its three main localities: Haqel, Hajoula, and Namoura.

It is one of three major Cretaceous lagerstätte in Lebanon, alongside the older (Barremian-aged) Lebanese amber and the younger (Santonian-aged) Sahel Alma site.

Description

It is primarily Cenomanian in age. The formation laterally varies from east to west; the western lowland "coastal" sequence is over 2000 metres thick and primarily consists of deep water limestone and chalk, while the eastern "mountain" sequence is 500–700 metres thick and consists of shallow water limestone.^[1]

Pterosaur fossils have been recovered from the formation, including those of Mimodactylus^[2] and Microtuban.^[3] The pythonomorphs Pontosaurus, Eupodophis and probably Aphanizocnemus are known from the formation.^[4] Insects are also known from compression fossils, including those of a dragonfly,^[5] enigmatic pond-skater like insect Chresmoda.^[6] and a mesoblattinid cockroach.^[7] Compression fossils of angiosperm Sapindopsis are also known.^[8] Numerous species of fossil fish are known,^[9] including the ionoscopid Spathiurus dorsalis^[10] and pycnodontid Flagellipinna rhomboides.^[11] An incredible diversity of fossil pycnodonts of various body plans is known, in addition to the earliest representatives of modern fish groups such as acanthomorphs and eels.^[12]

History

The Sannine Formation and its fossil fishes have been known of since Roman Phoenicia. The earliest mention of them is in the Chronicon by Eusebius around 314 CE, who mentions fossil fishes commonly being found during quarrying operations in the mountains, and cites them as evidence of the Biblical Flood. The site that these fish were recovered from is thought to correspond to the modern Haqel site. Around 1250, fossil fishes were presented to Louis IX during the Seventh Crusade and documented by Jean de Joinville, although it remains uncertain whether these originated from Haqel or the younger Sahel Alma site.^[12][13]

The first European to visit a Lebanese fossil fish locality in person was Cornelis de Bruijn in 1698, who illustrated a fossil fish that corresponds with the species Prionolepis cataphractus from Haqel. The first scientific presentation on the fossil fishes of Haqel was in 1703 at the French Academy of Sciences, and the first scientific studies were conducted by Henri Marie Ducrotay de Blainville in 1818. Lady Hester Stanhope collected and circulated fossil fishes from Haqel and Sahel Alma during the early 19th century, contributing to the study of these specimens by scientists such as Louis Agassiz. The Hadjoula site was only discovered in the late 19th century, and the Namoura site in the early 20th century.^[12][13]

Paleobiota

Jawless fish

Agnathans reported from the Sannine Formation
Genus	Species	Presence	Notes	Images
Tethymyxine	T. tapirostrum	Hâdjoula	One of the earliest known definitive fossil hagfish.[14]

Cartilaginous fish

Chondrichthyans reported from the Sannine Formation[15]
Genus	Species	Presence	Notes	Images
Cretodus	C. "longiplicatus"	Hâdjoula	A pseudoscapanorhynchid shark.
	C. semiplicatus	Haqel & Hâdjoula
Cyclobatis	C. major	Haqel & Hâdjoula	A cyclobatid ray.[16][17]
	C. oligodactylus	Haqel, Hâdjoula & Namoura[17]
	C. tuberculatus	Haqel
Libanopristis	L. hiram	Haqel & Hâdjoula	A sawskate.
Mesiteia	M. emiliae	Haqel	A bamboo shark.
Micropristis	M. solomonis	Hâdjoula	A sawskate.
Pararhinchodon	P. lehmani	Haqel	A collared carpet shark.
Pseudocorax	P. kindlimmani	Haqel	A false crow shark.[18]
Rajorhina	R. expansa	Hâdjoula	A skate.
Rhinobatos	R. grandis	Haqel	A guitarfish.
	R. hakelensis
	R. maronita
	R. whitfieldi	Hâdjoula
Rhombopterygia	R. rajoides	Hâdjoula	A guitarfish.
Scyliorhinus	S. arambourgi	Hâdjoula	A catshark.
	S. bloti	Haqel
Squalicorax	S. falcatus	Namoura	A crow shark.

Bony fish

Fish reported from the Sannine Formation[15]
Genus	Species	Presence	Notes	Images
Abisaadia	A. hakelensis	Haqel	An eel.
Abisaadichthys	A. libanicus	Hâdjoula	A protobramid tselfatiiform.
Acrognathus	A. dodgei	Haqel & Hâdjoula	A greeneye.
Acrorhinichthys	A. poyatoi	Haqel & Namoura	A pycnodont of uncertain affinities.
Aipichthys	A. minor	Haqel & Hâdjoula	A lamprimorph.
	A. oblongus	Haqel
	A. velifer	Haqel
Anguillavus	A. mazeni	Haqel & Hâdjoula	An eel.
	A. quadripinnis	Hâdjoula
Apatepopholis	A. laniatus	Haqel, Hâdjoula & Namoura	An apatepopholid aulopiform.
Aphanepygus	A. dorsalis	Haqel	An aphanepygid.
Akromystax	A. tilmachiton	Haqel & Namoura	A pycnodont of uncertain affinities.
Armigatus	A. alticorpus	Namoura[19] and Haqel[20]	A clupeomorph[20][19]
	A. brevissimus	Haqel and Hâdjoula[20]	An armigatid clupeomorph[20]
	A. namourensis	Namoura[19]	A clupeomorph[20]
Belonostomus	B. "sp. 1"	Namoura[19]	An aspidorhynchid. 2 unnamed species awaiting better material[19]
	B. "sp. 2"
Berycopsis	B. pulcher	Hâdjoula	A beardfish.
Capassopiscis	C. pankowskii	Haqel	A pantodontid.[21]
Charitopsis	C. spinosus	Haqel & Hâdjoula	A gonorynchid.
Coccodus	C. armatus	Haqel	A coccodontid pycnodont.
	C. insignis	Hâdjoula
Corusichthys	C. megacephalus	Haqel	A coccodontid pycnodont.
Cretapantodon	C. polli	Haqel	A pantodontid.[22]
Cryptoberyx	C. minimus	Hâdjoula	An indeterminate acanthopterygian.
Ctenodentelops	C. striatus	Namoura[19]	An elopid.[19]
Ctenothrissa	C. protodorsalis	Haqel	A ctenothrissiform acanthomorph.
	C. signifer	Hâdjoula
	C. vexillifer	Haqel
?Cylindracanthus	?C. libanicus	Haqel & Hâdjoula	An indeterminate ray-finned fish known only from rostral spines.
Davichthys	D. gardnieri	Haqel	An elopid.
Diplomystus	D. birdi	Haqel & Hâdjoula	An armigatid clupeomorph.
Ducrotayichthys	D. cornutus	Haqel	A gladiopycnodontid pycnodont.
Enchelion	E. montium	Haqel & Hâdjoula	An eel.
Enchodus	E. marchesetti	Haqel & Hâdjoula	An enchodontid aulopiform.
	E. mecoanalis	Namoura[19]
	E. "sp. 2."	Namoura
Eubiodectes	E. libanicus	Haqel, Hâdjoula & Namoura	A cladocyclid ichthyodectiform.
Eurypholis	E. boissieri	Haqel, Hâdjoula & Namoura	An enchodontid aulopiform.
Eusebichthys	E. byblosi	Haqel	A protobramid tselfatiiform.
	E. "sp. 1"	Haqel
	E. "sp. 2"	Hâdjoula
Exocoetoides	E. minor	Haqel & Hâdjoula	A cheirothricid.
Flagellipinna	F. rhomboides	Haqel	A pycnodontid pycnodont.[11]
Gebrayelichthys	G. uyenoi	Haqel	A gebrayelichthyid pycnodont.
	G. vexillarius	Haqel
Gaudryella	G. gaudryi	Haqel, Hâdjoula & Namoura	An indeterminate euteleost.
Gharbouria	G. libanica	Namoura	An indeterminate euteleost.
Gigapteryx	G. tethyestris	Namoura[19]	A lamprimorph.[19][23]
Gladiopycnodus	G. byrnei	Hajoula	A gladiopycnodontid pycnodont.[24]
	G. karami	Haqel
Ginsburgia	G. operta	Haqel & Hâdjoula	An indeterminate euteleost.
Hajulia	H. multidens	Haqel & Hâdjoula	A bonefish.
Hakelia	H. laticauda	Haqel & Hâdjoula	An indeterminate neoteleost.
Hakeliosomus	H. hakelensis	Haqel & Hâdjoula	A gonorynchid.
Haqelpycnodus	H. picteti	Haqel	A pycnodontid pycnodont.
Hastichthys	H. gracilis	Haqel & Namoura	A dercetid aulopiform.
Hayenchelys	H. germanus	Hâdjoula	An eel.
Hayolperichthys	H. pectospinus	Haqel	A gladiopycnodontid pycnodont.
Hemisaurida	H. hakelensis	Haqel	A halecid aulopiform.
Hensodon	H. spinosus	Haqel	A coccodontid pycnodont[25]
Heterothrissa	H. signeuxae	Haqel	A ctenothrissiform acanthomorph.
Hgulichthys	H. spinus	Hâdjoula	A trachichthyiform.
Humilichthys	H. orientalis	Hâdjoula	A ctenothrissiform acanthomorph.
Ichthyoceros	I. spinosus	Haqel	A coccodontid pyncodont.
Ichthyotringa	I. delicata	Haqel & Hâdjoula	An ichthyotringid aulopiform.
Joinvillichthys	J. kriweti	Haqel & Hâdjoula	A gladiopycnodontid pycnodont.
	J. lindstroemi	Haqel
Lebonichthys	L. lewisi	Hâdjoula	A bonefish.[19]
	L. namourensis	Namoura[19]
Lebrunichthys	L. nammourensis	Namoura	A pachyrhizodontid.
Libanoberyx	L. spinosus		A trachichthyiform.
Lissoberyx	L. arambourgi	Hâdjoula	A trachichthyiform.
	L. dayi
	L. denticulatus
Luenchelys	L. minimus	Namoura	An eel.
Macropomoides	M. orientalis	Haqel & Hâdjoula	A latimeriid coelacanth.
Maraldichthys	M. verticalis	Haqel	A gebrayelichthyid pycnodont.
Microcapros	M. libanicus	Hâdjoula	A quaesitoberycid beryciform.[26]
Monocerichthys	M. scheuchzeri	Haqel & Hâdjoula	A gladiopycnodontid pycnodont.
Nematonotus	N. bottae	Haqel & Namoura	A flagfin.
	N. longispinus	Hâdjoula
Nursallia	N. goedeli	Haqel, Hâdjoula & Namoura	A pycnodontid pycnodont.
Omosoma	O. sp.	Namoura	Possibly an indeterminate beardfish[26], potentially a new genus.
Ornategulum	O. sardinioides	Haqel & Namoura	A clupeomorph.
Palaeopantodon	P. vandersypeni	Haqel[27]	A pantodontid.
Pankowskichthys	P. libanicus	Haqel & Namoura	A gladiopycnodontid pycnodont.
Pankowskipiscis	P. haqelensis	Haqel[28]	A pantodontid.
Paracentrus	P. lebanonensis	Namoura[19]	A holocentriform[19]
Paracoccodus	P. woodwardi	Haqel	A coccodontid pyncodont.
Pateroperca	P. libanica	Hâdjoula	A ctenothrissiform acanthomorph.
	P. robusta
Pattersonichthys	P. delicatus	Hâdjoula	A ctenothrissiform acanthomorph.
Pattersonoberyx	P. pharsus	Haqel	A potential holocentriform.[26]
Petalopteryx	P. syriacus	Haqel	An aphanepygid.
Petersichthys	P. libanicus	Haqel	A pantodontid.
Pharmacichthys	P. numismalis	Haqel	A pharmacichthyid lamprimorph.
	P. venenifer
Phoenicolepis	P. arcuatus	Hâdjoula	A ctenothrissiform acanthomorph.
Phylactocephalus	P. microlepis	Haqel & Hâdjoula	A halecid aulopiform.
Plectocretacicus	P. clarae	Haqel	A potential tetraodontiform.
Plesioberyx	P. discoides	Haqel	An indeterminate acanthopterygian.
	P. maximus
Prognathoglossum	P. kalassyi	Haqel	A pantodontid.
Prionolepis	P. cataphractus	Haqel & Hâdjoula	A prionolepid aulopiform.
Protobrama	P. avus	Haqel & Hâdjoula	A protobramid tselfatiiform.
	P. woodwardi	Haqel
Pseudoberyx	P. bottae	Haqel & Hâdjoula	An indeterminate teleost.
	P. grandis	Haqel
	P. syriacus	Haqel & Hâdjoula
Pycnosteroides	P. levispinus	Hâdjoula	A pycnosteroidid lamprimorph.
Quaesitoberyx	Q. minutus	Hâdjoula	A quaesitoberycid beryciform.[26]
Rhamphoichthys	R. taxidiotis	Haqel[29]	A billfish-like plethodid tselfatiiform.
Rhinopycnodus	R. gabriellae	Haqel	A pycnodontid pycnodont.
Rhynchodercetis	R. hakelensis	Haqel & Hâdjoula	A dercetid aulopiform.
	R. serpentinus	Hâdjoula
Rostropycnodus	R. gayeti	Haqel	A gladiopycnodontid pycnodont.
Sardinioides	S.	Haqel & Hâdjoula	A sardinioidid myctophiform.
	S. minimus
	S. pontivagus
Scombroclupea	S. diminuta	Namoura[19]	A clupeomorph.[19]
	S. macrophthalma	Haqel & Hâdjoula
Scutatoclupea	S. bacchiai	Namoura	A paraclupeid clupeomorph.
Sedenhorstia	S. dayi	Hâdjoula	A tarpon.
	S. libanica	Haqel
	S. orientalis	Hâdjoula
Serrilepis	S. minor	Namoura[19]	A halecid aulopiform.[19]
	S. prymnostrigos
Sigmapycnodus	S. giganteus	Haqel	A pycnodontid pycnodont.
Sorbinichthys	S. elusivo	Namoura	A sorbinichthyid clupeomorph.
Spathiurus	S. dorsalis	Haqel	An ionoscopiform.[10]
Spinascutichthys	S. pankowskiae	Namoura	An enchodontoid aulopiform.[30]
Stanhopeichthys	S. libanicus	Hâdjoula	A pachyrhizodontid.
Stenoprotome	S. hamata	Haqel	A gladiopycnodontid pycnodont.
Stichocentrus	S. elegans	Hâdjoula	A holocentriform.
	S. liratus
	S. spinulosus
	S. sp.
Stichopteryx[26]	S. lewisi	Haqel & Hâdjoula	A trachichthyiform.
Telepholis	T. tenuis	Haqel & Hâdjoula	A cheirothricid.
Trewavasia	T. carinata	Haqel	A coccodontid pycnodont.[25]
Tricertichthys	T. wenzi	Haqel	A gladiopycnodontid pycnodont.
Triplomystus	T. noorae	Namoura[19]	A paraclupeid clupeomorph.[19]
	T. oligoscutatus
Volcichthys	V. sp.	Haqel	An indeterminate aulopiform or sardinioid myctophiform.
Ypsiloichthys	Y. sibelleae	Haqel[31]	A teleost.

Amniotes

Amniotes reported from the Sannine Formation
Genus	Species	Presence	Notes	Images
Aphanizocnemus	A. libanensis	Although the type locality is unknown, it is said to "almost certainly" originate from the Sannine Formation.	A marine lizard. [4]
Enantiophoenix	E. electrophyla	Namoura	An enantiornithine bird.[32]
Eupodophis	E. descouensi	Namoura	An early marine simoliophiid snake with hind limbs.[4]
Microtuban	M. altivolans	Hâdjoula	An azdarchoid pterosaur.[3]
Mimodactylus	M. libanensis	Hâdjoula	An istiodactyliform pterosaur.[2]
Pontosaurus	P. kornhuberi	Namoura	A marine dolichosaurid lizard.[4]
Rhinochelys	R. nammourensis	Adult and multiple hatchling remains known, suggesting a nesting site was nearby.[33]	A marine protostegid turtle.[33]

Invertebrates

Invertebrates reported from the Sannine Formation
Genus	Species	Presence	Notes	Images
Allocrioceras	A. cf. annulatum		An anisoceratid ammonite.[18][34]
Boreopeltis	B. smithi	Haqel or Hâdjoula	A plesioteuthid cephalopod.[35]
Callianassa	C. sp.	Hâdjoula	A callianassid crustacean.
Chresmoda	C. libanica	Namoura	A water strider-like chresmodid insect.[6]
Cretacalliax	C. levantina	Hâdjoula	A callianopsid ghost shrimp.[36]
Dorateuthis	D. syriaca	Sâhel Aalma	A plesioteuthid cephalopod.[35]
Globulacaris	G. garassinoi	Haqel	A protozoeid thylacocephalan.
Glyphea	G. damesi	Hâdjoula	A glypheoid crustacean.
Jasus	J. sp.	Hâdjoula	A spiny lobster.
Keuppia	K. levante	Hâdjoula	An early octopus. 2 or 3 species known from the same locality[37]
	K. hyperbolaris
	K. sp.
Lebanoaktassia	L. curiosa	Hâdjoula	An aktassiid dragonfly.[38]
Libanocordulia	L. debiei	Hâdjoula	A libanocorduliid dragonfly[5]
Libanogomphus	L. lionelcavini	Hâdjoula	A libanogomphid dragonfly.[39]
Libanoliupanshania	L. mimi	Hâdjoula	A liupanshaniid dragonfly[5]
Mantelliceras	M. mantelli	Haqel	"the occurrence of Mantelliceras mantelli is regarded doubtful, as this species has been used for various taxa within the family Acanthoceratidae de Grossouvre, 1894, and specimens have never been figured"[18][34]
Mieroblattina	M. pacis	Nammoura	A mesoblattinid cockroach.[7]
Palaeopalinurellus	P. jbeilensis	Haqel	A slipper lobster.[40]
Paradollocaris	P. vannieri	Hâdjoula	A dollocaridid thylacocephalan.
Phoenice	P. pasinii	Haqel	A boxer shrimp.
Styletoctopus	S. annae	Hâqel	An early octopus.[37]
Tachypleus	T. syriacus	Hâdjoula	A horsehoe crab.[41]
Thylacocaris	T. schrami	Hâdjoula	A dollocaridid thylacocephalan.

Plants

See also

• List of pterosaur-bearing stratigraphic units
• Paleontology in Lebanon

References