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Sannine Formation

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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 from the Tethys Ocean within its three main localities: Haqel (alternatively Hakel or Haqil), Hjoula (alternatively Hadjoula, Hajoula, or Hgula), and Nammoura (alternatively Namoura).[1]

Sannine Formation
Stratigraphic range: Late Cenomanian
~95–93.5 Ma
↓
Fossil block from Haqel with fishes and a lobster
TypeGeological formation
UnderliesMaameltain Formation
OverliesHammana Formation
ThicknessCoastal: >2,000 m (6,600 ft)
Mountains: 500–700 m (1,600–2,300 ft)
Lithology
PrimaryChalk, limestone
OtherMarl
Location
Coordinates34°06′N 35°42′E / 34.1°N 35.7°E / 34.1; 35.7
Approximate paleocoordinates12°54′N 30°24′E / 12.9°N 30.4°E / 12.9; 30.4
RegionJabal Lubnan
Country Lebanon
Type section
Named forMount Sannine
Sannine Formation is located in Lebanon
Sannine Formation
Sannine Formation (Lebanon)

It is one of three major Cretaceous lagerstätte in Lebanon, alongside the older (Barremian-aged) Lebanese amber and the younger (Santonian-aged) Sahel Aalma site. The Sannine Formation localities, combined with Sahel Alma, are together referred to as the "Fish Beds" of Lebanon.[1]

Description

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It is primarily Cenomanian in age, with Haqel and Hjoula being late Cenomanian, while the slightly older Nammoura site is middle Cenomanian.[1] Although Lebanon is now a part of Asia, the depositional environment for both formations would have been located off the coast of northern Africa during the Cretaceous, which would technically make their biota African rather than Asian.[2][3]

The Cretaceous formations of Lebanon laterally vary from east to west; the western lowland "coastal" sequence (containing Sahel Aalma) is over 2000 metres thick and primarily consists of deep water limestone and chalk, while the eastern "mountain" sequence (containing the Sannine Formation) is 500–700 metres thick and consists of shallow water limestone.[4] Two different shallow-water environments are preserved by the formation: Nammoura preserves a nearshore inner continental shelf environment with significant terrestrial input, while Haqel & Hjoula preserve an outer continental shelf environment. The neritic nature of these habitats contrasts with the other major marine lagerstätte of the region, Sahel Aalma, which is thought to have been deposited in a deepwater continental slope environment.[1]

 
Modern position of the Hjoula locality compared to its Cretaceous location.

Pterosaur fossils have been recovered from the formation, including those of Mimodactylus[3] and Microtuban.[5] The pythonomorphs Pontosaurus, Eupodophis and probably Aphanizocnemus are known from the formation.[6] Insects are also known from compression fossils, including those of a dragonfly,[7] enigmatic pond-skater like insect Chresmoda.[8] and a mesoblattinid cockroach.[9] Compression fossils of angiosperm Sapindopsis are also known.[10] Numerous species of fossil fish are known,[11] including the ionoscopid Spathiurus dorsalis[12] and pycnodontid Flagellipinna rhomboides.[13] An incredible diversity of fossil pycnodonts of various, highly unusual body plans is known, in addition to the earliest representatives of modern fish groups such as acanthomorphs, African butterflyfish and eels.[1]

History

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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 (references to earlier mentions by Herodotus are thought to be apocryphal), 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.[1][14]

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 Nammoura site in the early 20th century.[1][14]

Paleobiota

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Vertebrates

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Jawless fish

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Agnathans reported from the Sannine Formation
Genus Species Presence Notes Images
Tethymyxine T. tapirostrum Hjoula One of the earliest known definitive fossil hagfish.[15]  

Cartilaginous fish

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Chondrichthyans reported from the Sannine Formation[16][17]
Genus Species Presence Notes Images
Cantioscyllium C. decipiens Haqel & Hjoula A nurse shark.
C. sp. Hjoula
Crassescyliorhinus C. cf. germanicus Hjoula A catshark.
Cretalamna C. sp. Hjoula An otodontid shark, formerly identified as C. appendiculata.[18]  
Cretodus C. "longiplicatus" Hjoula A pseudoscapanorhynchid shark.
C. semiplicatus Haqel & Hjoula  
Cretoxyrhina C. mantelli Hjoula A ginsu shark.  
Cyclobatis C. major Haqel & Hjoula A cyclobatid ray.[19][20]  
C. oligodactylus Haqel, Hjoula & Nammoura[20]  
C. tuberculatus Haqel
Deania D. sp. Haqel A gulper shark.
 
Ginglymostomatidae indet. Hjoula A nurse shark.
Haimirichia H. amonensis Haqel & Nammoura A haimirichiid mackerel shark.
Hemiscylliidae indet. Haqel & Hjoula A bamboo shark.
Heterodontus H. sp. Hjoula A bullhead shark.  
Libanopristis L. hiram Haqel & Hjoula A sawskate.  
Mesiteia M. emiliae Haqel A mesiteiid carpet shark.
Micropristis M. solomonis Hjoula A sawskate.  
Odontaspididae indet. Hjoula A sand shark.  
Odontaspis O. aculeatus Hjoula A sand shark.  
O. sp.
Parascyllidae indet. Haqel & Hjoula A collared carpet shark.
Paranomotodon P. sp. Hjoula A thresher shark.
Protoscyliorhinus P. sp. Haqel A catshark.
Pseudocorax P. kindlimmani Haqel A false crow shark.[21]  
Rajorhina R. expansa Hjoula A skate.  
'Rhinobatos' (gen. nov. 1) R. grandis Haqel A potential relative of wedgefishes. Likely its own genus.[22]
R. maronita  
'Rhinobatos' (gen. nov. 3) R. hakelensis Haqel A guitarfish-like ray. Likely its own genus, and its status as a guitarfish is disputed.[22]  
R. whitfieldi Hjoula  
Rhombopterygia R. rajoides Hjoula A guitarfish.  
Scyliorhinidae indet. Haqel A catshark.
Scyliorhinus S. arambourgi Hjoula A catshark.  
S. bloti Haqel
S. sp.
Squalicorax S. falcatus Nammoura A crow shark.  
S. sp. Haqel, Hjoula & Nammoura
Squatina S. sp. Haqel An angelshark.  
Triakidae indet. Haqel A houndshark.

Ray-finned fish

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Actinopterygians reported from the Sannine Formation[16]
Genus Species Presence Notes Images
Abisaadia A. hakelensis Haqel An eel.
Abisaadichthys A. libanicus Hjoula A protobramid tselfatiiform.
Acrognathus A. dodgei Haqel & Hjoula A greeneye.
Acrorhinichthys A. poyatoi Haqel & Nammoura A pycnodont of uncertain affinities.
Aipichthys A. minor Haqel & Hjoula A lamprimorph.  
A. oblongus Haqel
A. velifer Haqel  
Anguillavus A. mazeni Haqel & Hjoula An eel.  
A. quadripinnis Hjoula
Apateopholidae indet. nov. gen. nov. sp. Haqel An apateopholid aulopiform.
Apateopholis A. laniatus Haqel, Hjoula & Nammoura An apateopholid aulopiform.
A. nov. sp. Nammoura
Aphanepygus A. dorsalis Haqel An aphanepygid.  
Akromystax A. tilmachiton Haqel & Nammoura A pycnodont of uncertain affinities.
Armigatus A. alticorpus Nammoura[23] and Haqel[24] A clupeomorph[24][23]
A. brevissimus Haqel and Hjoula[24] An armigatid clupeomorph[24]
 
A. namourensis Nammoura[23] A clupeomorph[24]
Belonostomus B. "sp. 1" Nammoura[23] An aspidorhynchid. 2 unnamed species awaiting better material[23]  
B. "sp. 2"
Benthesikyme B. serpentina Haqel & Hjoula A dercetid aulopiform.[25]  
Berycopsis B. pulcher Hjoula A beardfish.  
Capassopiscis C. pankowskii Haqel A pantodontid.[26]
Charitopsis C. spinosus Haqel & Hjoula A gonorynchid.  
Chirocentrites C. sp. Nammoura A cladocyclid ichthyodectiform.  
Coccodus C. armatus Haqel A coccodontid pycnodont.
C. insignis Hjoula  
Corusichthys C. megacephalus Haqel A coccodontid pycnodont.  
Cretapantodon C. polli Haqel A pantodontid.[27]
Cryptoberyx C. minimus Hjoula An indeterminate acanthopterygian.
Ctenodentelops C. striatus Nammoura[23] An elopid.[23]
Ctenothrissa C. protodorsalis Haqel A ctenothrissiform acanthomorph.
C. signifer Hjoula
C. vexillifer Haqel  
?Cylindracanthus ?C. libanicus Haqel & Hjoula An indeterminate ray-finned fish known only from rostral spines.
Davichthys D. gardnieri Haqel An elopid.  
Dercetidae indet. nov. gen. nov. sp. Hjoula A dercetid aulopiform.
Diplomystus D. birdi Haqel & Hjoula An armigatid clupeomorph.  
Ducrotayichthys D. cornutus Haqel A gladiopycnodontid pycnodont.
Enchelion E. montium Haqel & Hjoula An eel.  
Enchodus E. marchesetti Haqel & Hjoula An enchodontid aulopiform.  
E. mecoanalis Nammoura[23]
E. "sp. 1." Haqel
E. "sp. 2." Nammoura
Eubiodectes E. libanicus Haqel, Hjoula & Nammoura A cladocyclid ichthyodectiform.  
Eurypholidae indet. nov. gen. nov. sp. Nammoura An enchodontid aulopiform.
Eurypholis E. boissieri Haqel, Hjoula & Nammoura An enchodontid aulopiform.  
Eusebichthys E. byblosi Haqel A protobramid tselfatiiform.
E. "sp. 1" Haqel
E. "sp. 2" Hjoula
Exocoetoides E. minor Haqel & Hjoula A cheirothricid.  
Flagellipinna F. rhomboides Haqel A pycnodontid pycnodont.[13]
Gebrayelichthys G. uyenoi Haqel A gebrayelichthyid pycnodont.  
G. vexillarius Haqel
Gaudryella G. gaudryi Haqel, Hjoula & Nammoura An indeterminate euteleost.  
Gharbouria G. libanica Nammoura An indeterminate euteleost.
Gigapteryx G. tethyestris Nammoura[23] A lamprimorph.[23][28]
Gladiopycnodus G. byrnei Hjoula A gladiopycnodontid pycnodont.[29]
G. karami Haqel
Ginsburgia G. operta Haqel & Hjoula An indeterminate euteleost.
Hajulia H. multidens Haqel & Hjoula A bonefish.
Hakelia H. laticauda Haqel & Hjoula An indeterminate neoteleost.
Hakeliosomus H. hakelensis Haqel & Hjoula A gonorynchid.  
Haqelpycnodus H. picteti Haqel A pycnodontid pycnodont.
 
Hastichthys H. gracilis Haqel & Nammoura A dercetid aulopiform.
Hayenchelys H. germanus Hjoula An eel.  
Hayolperichthys H. pectospinus Haqel A gladiopycnodontid pycnodont.
Hemisaurida H. hakelensis Haqel A halecid aulopiform.  
Hensodon H. spinosus Haqel A coccodontid pycnodont[30]  
Heterothrissa H. signeuxae Haqel A ctenothrissiform acanthomorph.
Hgulichthys H. spinus Hjoula A trachichthyiform.
Humilichthys H. orientalis Hjoula A ctenothrissiform acanthomorph.
Ichthyoceros I. spinosus Haqel A coccodontid pyncodont.  
Ichthyotringa I. delicata Haqel & Hjoula An ichthyotringid aulopiform.  
Joinvillichthys J. kriweti Haqel & Hjoula A gladiopycnodontid pycnodont.
J. lindstroemi Haqel
Lebonichthys L. lewisi Hjoula A bonefish.[23]
L. namourensis Nammoura[23]
Lebrunichthys L. nammourensis Nammoura A pachyrhizodontid.
Libanoberyx L. spinosus A trachichthyiform.
Lissoberyx L. arambourgi Hjoula A trachichthyiform.  
L. dayi
L. denticulatus
Luenchelys L. minimus Nammoura An eel.
Maraldichthys M. verticalis Haqel A gebrayelichthyid pycnodont.  
Microcapros M. libanicus Hjoula A quaesitoberycid beryciform.[31]
Monocerichthys M. scheuchzeri Haqel & Hjoula A gladiopycnodontid pycnodont.
Nematonotus N. bottae Haqel & Nammoura A flagfin.  
N. longispinus Hjoula  
'Nursallia' 'N.' goedeli Haqel, Hjoula & Nammoura A pycnodontid pycnodont.  
'N.' tethysensis Nammoura and Hjoula
Omosoma O. sp./gen. nov. sp. nov. Nammoura Possibly an indeterminate beardfish,[31] potentially a new genus.
Ornategulum O. sardinioides Haqel & Nammoura A clupeomorph.
Palaeopantodon P. vandersypeni Haqel[32] A pantodontid.
Pankowskichthys P. libanicus Haqel & Nammoura A gladiopycnodontid pycnodont.
Pankowskipiscis P. haqelensis Haqel[33] A pantodontid.
Paracentrus P. lebanonensis Nammoura[23] A holocentriform[23]
P. sp. nov.
Paraclupeidae indet. nov. gen. 1 Nammoura A paraclupeid clupeomorph; two different genera.
nov. gen. 2. Hjoula
Paracoccodus P. woodwardi Haqel A coccodontid pyncodont.
Pateroperca P. libanica Hjoula A ctenothrissiform acanthomorph.
P. robusta
Pattersonichthys P. delicatus Hjoula A ctenothrissiform acanthomorph.
Pattersonoberyx P. pharsus Haqel A potential holocentriform.[31]
Petalopteryx P. syriacus Haqel An aphanepygid.
Petersichthys P. libanicus Haqel A pantodontid.
Pharmacichthys P. numismalis Haqel A pharmacichthyid lamprimorph.
P. venenifer  
Phoenicolepis P. arcuatus Hjoula A ctenothrissiform acanthomorph.
Phylactocephalus P. microlepis Haqel & Hjoula 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 & Hjoula A prionolepid aulopiform.  
Protobrama P. avus Haqel & Hjoula A protobramid tselfatiiform.  
P. woodwardi Haqel
Protobramidae indet. nov. gen. nov. sp. Nammoura A protobramid tselfatiiform.
Pseudoberyx P. bottae Haqel & Hjoula An indeterminate teleost.
P. grandis Haqel
P. syriacus Haqel & Hjoula
Pycnosteroides P. levispinus Hjoula A pycnosteroidid lamprimorph.
Quaesitoberyx Q. minutus Hjoula A quaesitoberycid beryciform.[31]
Rhamphoichthys R. taxidiotis Haqel[34] A billfish-like plethodid tselfatiiform.  
Rhinopycnodus R. gabriellae Haqel A pycnodontid pycnodont.
Rhynchodercetis R. hakelensis Haqel & Hjoula A dercetid aulopiform.  
Rostropycnodus R. gayeti Haqel A gladiopycnodontid pycnodont.
Sardinioides S. attenuatus Haqel & Hjoula A sardinioidid myctophiform.  
S. minimus
S. pontivagus
Saurorhamphus S. giorgiae Nammoura An enchodontid aulopiform.
Scombroclupea S. diminuta Nammoura[23] A clupeomorph.[23]  
S. macrophthalma Haqel & Hjoula
Scutatoclupea S. bacchiai Nammoura A paraclupeid clupeomorph.
Sedenhorstia S. dayi Hjoula A tarpon.  
S. libanica Haqel
S. orientalis Hjoula
Serrilepis S. minor Nammoura[23] A halecid aulopiform.[23]
S. prymnostrigos
Sigmapycnodus S. giganteus Haqel A pycnodontid pycnodont.
Sorbinichthys S. elusivo Nammoura A sorbinichthyid clupeomorph.  
Spathiurus S. dorsalis Haqel An ionoscopiform.[12]
Spinascutichthys S. pankowskiae Nammoura An enchodontoid aulopiform.[35]
Stanhopeichthys S. libanicus Hjoula A pachyrhizodontid.
Stenoprotome S. hamata Haqel A gladiopycnodontid pycnodont.
Stichocentrus S. elegans Hjoula A holocentriform.  
S. liratus
S. spinulosus
S. sp.
Stichopteryx[31] S. lewisi Haqel & Hjoula A trachichthyiform.
Telepholis T. tenuis Haqel & Hjoula A cheirothricid.
Trewavasia T. carinata Haqel A coccodontid pycnodont.[30]  
Tricerichthys T. wenzi Haqel A gladiopycnodontid pycnodont.
Triplomystus T. noorae Nammoura[23] A paraclupeid clupeomorph.[23]
T. oligoscutatus
Volcichthys V. sp. Haqel An indeterminate aulopiform or sardinioid myctophiform.
Ypsiloichthys Y. sibelleae Haqel[36] A teleost of uncertain affinities.  

Lobe-finned fish

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Sarcopterygians reported from the Sannine Formation
Genus Species Presence Notes Images
Macropomoides M. orientalis Haqel & Hjoula A latimeriid coelacanth.  

Amniotes

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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.[6]
 
Enantiophoenix E. electrophyla Nammoura An enantiornithine bird.[37]
Eupodophis E. descouensi Nammoura An early marine simoliophiid snake with hind limbs.[6]
 
Microtuban M. altivolans Hjoula An azdarchoid pterosaur.[5]  
Mimodactylus M. libanensis Hjoula An istiodactyliform pterosaur.[3]
 
Pontosaurus P. kornhuberi Nammoura A marine dolichosaurid lizard.[6]
 
Rhinochelys R. nammourensis Nammoura A marine protostegid turtle. Adult and multiple hatchling remains known, suggesting a nesting site was nearby.[38]
 
Theropoda indet. Nammoura The partial arm of a theropod dinosaur, the only non-avialan dinosaur material known from the formation. Now lost.[1]

Invertebrates

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Arthropods

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Arthropods reported from the Sannine Formation[39]
Genus Species Presence Notes Images
Acanthochirana A. smithwoodwardi Haqel & Hjoula An aegerid prawn.  
Acanthophoenicides A. peterpani A slipper lobster.
Aeger A. libanensis Haqel & Hjoula An aegerid prawn.  
Armacrangon A. edwardsi A glyphocrangonid shrimp.
?Bentheogennema B. phoenicia A benthesicymid crustacean.
Callianassa C. sp. Hjoula A callianassid crustacean.[40]
Calliaxina C. hadjoulaensis Hjoula A callianassid crustacean.
Carinacaris C. antennae A penaeid prawn.
C. teruzzi Haqel & Hjoula
Carpopenaeus C. callirostris Haqel & Hjoula A carpopenaeid prawn.  
C. septemspinatus Haqel & Hjoula  
Charbelicaris C. maronites Hjoula A slipper lobster.[41]
Chresmoda C. libanica Nammoura A water strider-like chresmodid insect.[8]  
Corazzatocarcinus C. hadjoulae Hjoula A necrocarcinid crab.[42]  
Cretaxiopsis C. libanotica An axiid crustacean.
Cretacalliax C. levantina Hjoula A callianopsid ghost shrimp.[43]
Epipenaeus E. abisaadorum An carpopenaeid shrimp.
Eryma E. oscari Hjoula An erymid "lobster".  
Gladicrangon G. coriacea A glyphocrangonid shrimp.
Globulacaris G. garassinoi Haqel A protozoeid thylacocephalan.
Glyphea G. damesi Hjoula A glypheoid crustacean.[40]  
Hakelocaris H. vavassorii Haqel & Hjoula A penaeid prawn.  
Homolopsis H. aff. edwardsii Hjoula A carrier crab.[42]
Jasus J. sp. Hjoula A spiny lobster.[40]  
Joinvilleicaris J. longirostris A penaeid prawn.
Libanoaxius L. beatae An axiid crustacean.
Libanocaris L. curvirostra A penaeid prawn.
L. rogeri Haqel & Hjoula
Lebanoaktassia L. curiosa Hjoula An aktassiid dragonfly.[44]
Libanocordulia L. debiei Hjoula A libanocorduliid dragonfly[7]
Libanogomphus L. lionelcavini Hjoula A libanogomphid dragonfly.[45]
Libanoliupanshania L. mimi Hjoula A liupanshaniid dragonfly[7]
Linuparus L. secretanae A spiny lobster.  
Macropenaeus M. incertus Haqel & Hjoula A penaeid prawn.  
Magila M. cretacica An axiid crustacean.
Microchela M. rostrata Haqel & Hjoula A penaeid prawn.
Mieroblattina M. pacis Nammoura A mesoblattinid cockroach.[9]
Notahomarus N. hakelensis Haqel & Hjoula A true lobster.  
Odontochelion O. bicarinatum A crangonid shrimp.
O. cretaceum Haqel & Hjoula
Palaeobenthonectes P. arambourgi A benthesicymid crustacean.
Palaeopalinurellus P. jbeilensis Haqel A slipper lobster.[46]
Palaeopandalus P. komaii A pandalid shrimp.
Palibacus P. praecursor Haqel & Hjoula A slipper lobster.  
Palinurus P. teruzzii A spiny lobster.  
Paracancrinos P. libanensis Hjoula A slipper lobster.[41]
Paradollocaris P. vannieri Hjoula A dollocaridid thylacocephalan.
Paranecrocarcinus P. cf. milbournei Hjoula A necrocarcinid crab. Specimen consumed by a catshark.[47]
Phalangiopsis P. antoinei A chirostylid squat lobster.
P. rogeri
Phoenice P. pasinii Haqel A boxer shrimp.[40]
Pseudastacus P. dubertreti Haqel & Hjoula A stenochirid "lobster".  
Pustulina P. cretacea[48] Haqel & Hjoula An erymid "lobster".  
Striadiogenes S. frigerioi Hjoula A diogenid hermit crab.[49]
Tachypleus T. syriacus Hjoula A horseshoe crab.[50]  
Telamonocarcinus T. gambulatus Haqel & Hjoula A dorippid crab.[42]
Thylacocaris T. schrami Hjoula A dollocaridid thylacocephalan.

Molluscs

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Molluscs reported from the Sannine Formation
Genus Species Presence Notes Images
Allocrioceras A. cf. annulatum An anisoceratid ammonite.[21][51]  
Boreopeltis B. smithi Haqel or Hjoula A plesioteuthid coleoid.[52]
Dorateuthis D. syriaca Haqel & Hjoula A plesioteuthid coleoid.[53]  
Edmunditeuthis E. bacchiai Haqel & Hjoula A trachyteuthid coleoid.[54][55]
Glyphidopsis G. waagei Haqel A trachyteuthid coleoid.[52]
Glyphiteuthis G. abisaadorum Haqel & Hjoula A trachyteuthid coleoid.[52][56]
G. freijii Haqel, Hjoula & Nammoura
G. libanotica Haqel & Hjoula
Keuppia K. levante Hjoula An early octopus. 2 or 3 species known from the same locality[57]
 
K. hyperbolaris
K. sp.
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"[21][51]  
Rachiteuthis R. acutali Hjoula A palaeololiginid coleoid.[58]
R. donovani Hâqel
Styletoctopus S. annae Hâqel An early octopus.[57]
Teudopsinia T. haasi Haqel A teudopsid coleoid.[54]
Syrionautilus S. libanoticus Hjoula A cymatoceratid nautiloid.[59]

Echinoderms

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Echinoderms reported from the Sannine Formation[60]
Genus Species Presence Notes Images
Antedon A. pinnulata Haqel A crinoid.  
Aspidochirotida indet. Hjoula A sea cucumber.[61]
Geocoma G. libanotica Haqel A brittle star.  

Annelids

edit
Annelids reported from the Sannine Formation[62]
Genus Species Presence Notes Images
Didone D. pulcherrima A didonid bristleworm.
Eunicites E. diopatroides A eunicid bristleworm.
E. falcatus
E. joinvillei
E. mariacristinae
Ferragutia F. cenomaniana A bristleworm.
Lumbriconereites L. garassinoi A bristleworm.
L. hadjulae
Palaeoaphrodite P. libanica An aphroditid bristleworm.
Phoeniciarabella P. caesaris A lumbrinerid bristleworm.
P. orensanzi
P. pinnulata
Teruzzia T. gryphoeides A lumbrinerid bristleworm.
T. pezzolii
T. pusilla
T. sagittifera
Rollinschaeta R. myoplena Haqel & Hjoula A fireworm.[63]

Plants

edit
Plants reported from the Sannine Formation
Genus Species Presence Notes Images
Aryskumia cf. A. zelkovifolia Nammoura Dicot known from a poorly preserved leaf with ovate blade.[64]
Nammouria N. gracilis An aquatic fern of uncertain affinities.[64]
Nammourophyllum N. altingioides A dicot known from leaves, a possible member of the Altingiaceae.[64]
Nupharanthus N. cretacea A dicot known by a preserved flower, possibly a member of the Nympheales.[64]
Parvileguminophyllum P. sp. Dicot known from a complete leaflet, possibly a member of the Fabaceae.[64]
'Phragmites' P. sp. A monocot of uncertain affinities, vaguely similar to the modern reed.[64]
Platycaryeae gen. et sp. indet. Infructescence of bracteate fruits resembling Platycarya americana.[64]
Pseudolarix P. sp. A conifer related to the modern golden larch.[64]  
Pseudotorellia P. sp. A gymnosperm of uncertain affinities.[64]
Pteris P. sp. A pteridacean fern.[64]  
Sapindopsis S. anhouryi A platanacean, related to the modern plane trees.[64]  
S. libanensis

See also

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References

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