Sannine Formation
Sannine Formation | |
---|---|
Stratigraphic range: Late Cenomanian ~95–93.5 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓ | |
![]() Fossil block from Haqel with fishes and a lobster | |
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 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]
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
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]
![](http://upload.wikimedia.org/wikipedia/commons/thumb/d/de/Mimodactylus_map.png/220px-Mimodactylus_map.png)
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
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
Vertebrates
Jawless fish
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
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
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 apatepopholid aulopiform. | |
Apatepopholis | A. laniatus | Haqel, Hjoula & Nammoura | An apatepopholid 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
Sarcopterygians reported from the Sannine Formation | ||||
---|---|---|---|---|
Genus | Species | Presence | Notes | Images |
Macropomoides | M. orientalis | Haqel & Hjoula | A latimeriid coelacanth. | ![]() |
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. [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 dinosaur material known from the formation. Now lost.[1] |
Invertebrates
Arthropods
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 horsehoe crab.[50] | ![]() |
Telamonocarcinus | T. gambulatus | Haqel & Hjoula | A dorippid crab.[42] | |
Thylacocaris | T. schrami | Hjoula | A dollocaridid thylacocephalan. |
Molluscs
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
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
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
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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{{cite journal}}
: CS1 maint: multiple names: authors list (link) - ^ Jattiot, Romain; Brayard, Arnaud; Fara, Emmanuel; Charbonnier, Sylvain (January 2015). "Gladius-bearing coleoids from the Upper Cretaceous Lebanese Lagerstätten: diversity, morphology, and phylogenetic implications". Journal of Paleontology. 89 (1): 148–167. Bibcode:2015JPal...89..148J. doi:10.1017/jpa.2014.13. ISSN 0022-3360.
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