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).

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.

Description

It is primarily Cenomanian in age, with Haqel and Hjoula being late Cenomanian, while the slightly older Nammoura site is middle Cenomanian. 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.

Map of Lebanon showing the locations of the 4 famous Upper Cretaceous Lagerstätten

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. 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.

Pterosaur fossils have been recovered from the formation, including those of Mimodactylus and Microtuban. The pythonomorphs Pontosaurus, Eupodophis and probably Aphanizocnemus are known from the formation. Insects are also known from compression fossils, including those of a dragonfly, enigmatic pond-skater like insect Chresmoda. and a mesoblattinid cockroach. Compression fossils of angiosperm Sapindopsis are also known. Numerous species of fossil fish are known, including the ionoscopid Spathiurus dorsalis and pycnodontid Flagellipinna rhomboides. 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.

History

Diagram of the depositional environments of the Upper Cretaceous Sannine Formation Lagerstätten with a selection of iconic flora and fauna

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.

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.

Paleobiota

Vertebrates

Jawless fish

Agnathans reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
TethymyxineT. tapirostrumHjoulaOne of the earliest known definitive fossil hagfish.

Cartilaginous fish

Chondrichthyans reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
CantioscylliumC. decipiensHaqel & HjoulaA nurse shark.
C. sp.Hjoula
CrassescyliorhinusC. cf. germanicusHjoulaA catshark.
CretalamnaC. sp.HjoulaAn otodontid shark, formerly identified as C. appendiculata.
CretodusC. "longiplicatus"HjoulaA pseudoscapanorhynchid shark.
C. semiplicatusHaqel & Hjoula
CretoxyrhinaC. mantelliHjoulaA Cretoxyrhinid.
CyclobatisC. majorHaqel & HjoulaA cyclobatid ray.
C. oligodactylusHaqel, Hjoula & Nammoura
C. tuberculatusHaqel
DeaniaD. sp.HaqelA gulper shark.
Ginglymostomatidae indet.HjoulaA nurse shark.
HaimirichiaH. amonensisHaqel & NammouraA haimirichiid mackerel shark.
Hemiscylliidae indet.Haqel & HjoulaA bamboo shark.
HeterodontusH. sp.HjoulaA bullhead shark.
LibanopristisL. hiramHaqel & HjoulaA sawskate.
MesiteiaM. emiliaeHaqelA mesiteiid carpet shark.
MicropristisM. solomonisHjoulaA sawskate.
Odontaspididae indet.HjoulaA sand shark.
OdontaspisO. aculeatusHjoulaA sand shark.
O. sp.
ParanomotodonP. sp.HjoulaA thresher shark.
PararhincodonP. lehmaniHaqelA collared carpet shark.
P. sp.Haqel & Hjoula
ProtoscyliorhinusP. sp.HaqelA catshark.
PseudocoraxP. kindlimmaniHaqelA false crow shark.
RajorhinaR. expansaHjoulaA skate.
'Rhinobatos' (gen. nov. 1)R. grandisHaqelA potential relative of wedgefishes. Likely its own genus.
R. maronita
'Rhinobatos' (gen. nov. 3)R. hakelensisHaqelA guitarfish-like ray. Likely its own genus, and its status as a guitarfish is disputed.
R. whitfieldiHjoula
RhombopterygiaR. rajoidesHjoulaA guitarfish.
Scyliorhinidae indet.HaqelA catshark.
ScyliorhinusS. arambourgiHjoulaA catshark.
S. blotiHaqel
S. sp.
SqualicoraxS. falcatusNammouraA crow shark.
S. sp.Haqel, Hjoula & Nammoura
SquatinaS. sp.HaqelAn angelshark.
Triakidae indet.HaqelA houndshark.

Ray-finned fish

Actinopterygians reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
AbisaadiaA. hakelensisHaqelAn eel.
AbisaadichthysA. libanicusHjoulaA protobramid tselfatiiform.
AcrognathusA. dodgeiHaqel & HjoulaA greeneye.
AcrorhinichthysA. poyatoiHaqel & NammouraA pycnodont of uncertain affinities.
AipichthysA. minorHaqel & HjoulaA lamprimorph.
A. oblongusHaqel
A. veliferHaqel
AnguillavusA. mazeniHaqel & HjoulaAn eel.
A. quadripinnisHjoula
Apateopholidae indet.nov. gen. nov. sp.HaqelAn apateopholid aulopiform.
ApateopholisA. laniatusHaqel, Hjoula & NammouraAn apateopholid aulopiform.
A. nov. sp.Nammoura
AphanepygusA. dorsalisHaqelAn aphanepygid.
AkromystaxA. tilmachitonHaqel & NammouraA pycnodont of uncertain affinities.
ArmigatusA. alticorpusNammoura and HaqelA clupeomorph
A. brevissimusHaqel and HjoulaAn armigatid clupeomorph
A. namourensisNammouraA clupeomorph
BelonostomusB. "sp. 1"NammouraAn aspidorhynchid. 2 unnamed species awaiting better material
B. "sp. 2"
BenthesikymeB. serpentinaHaqel & HjoulaA dercetid aulopiform.
BerycopsisB. pulcherHjoulaA beardfish.
CapassopiscisC. pankowskiiHaqelA pantodontid.
CharitopsisC. spinosusHaqel & HjoulaA gonorynchid.
ChirocentritesC. sp.NammouraA cladocyclid ichthyodectiform.
CoccodusC. armatusHaqelA coccodontid pycnodont.
C. insignisHjoula
CorusichthysC. megacephalusHaqelA coccodontid pycnodont.
CretapantodonC. polliHaqelA pantodontid.
CryptoberyxC. minimusHjoulaAn indeterminate acanthopterygian.
CtenodentelopsC. striatusNammouraAn elopid.
CtenothrissaC. protodorsalisHaqelA ctenothrissiform acanthomorph.
C. signiferHjoula
C. vexilliferHaqel
?Cylindracanthus?C. libanicusHaqel & HjoulaAn indeterminate ray-finned fish known only from rostral spines.
DavichthysD. gardnieriHaqelAn elopid.
Dercetidae indet.nov. gen. nov. sp.HjoulaA dercetid aulopiform.
DiplomystusD. birdiHaqel & HjoulaAn armigatid clupeomorph.
DucrotayichthysD. cornutusHaqelA gladiopycnodontid pycnodont.
EnchelionE. montiumHaqel & HjoulaAn eel.
EnchodusE. marchesettiHaqel & HjoulaAn enchodontid aulopiform.
E. mecoanalisNammoura
E. "sp. 1."Haqel
E. "sp. 2."Nammoura
EubiodectesE. libanicusHaqel, Hjoula & NammouraA cladocyclid ichthyodectiform.
Eurypholidae indet.nov. gen. nov. sp.NammouraAn enchodontid aulopiform.
EurypholisE. boissieriHaqel, Hjoula & NammouraAn enchodontid aulopiform.
EusebichthysE. byblosiHaqelA protobramid tselfatiiform.
E. "sp. 1"Haqel
E. "sp. 2"Hjoula
ExocoetoidesE. minorHaqel & HjoulaA cheirothricid.
FlagellipinnaF. rhomboidesHaqelA pycnodontid pycnodont.
FreigichthysF. elleipsisHjoulaA lamprimorph.
GebrayelichthysG. uyenoiHaqelA gebrayelichthyid pycnodont.
G. vexillariusHaqel
GaudryellaG. gaudryiHaqel, Hjoula & NammouraAn indeterminate euteleost.
GharbouriaG. libanicaNammouraAn indeterminate euteleost.
GigapteryxG. tethyestrisNammouraA lamprimorph.
GladiopycnodusG. byrneiHjoulaA gladiopycnodontid pycnodont.
G. karamiHaqel
GinsburgiaG. opertaHaqel & HjoulaAn indeterminate euteleost.
HajuliaH. multidensHaqel & HjoulaA bonefish.
HakeliaH. laticaudaHaqel & HjoulaAn indeterminate neoteleost.
HakeliosomusH. hakelensisHaqel & HjoulaA gonorynchid.
HaqelpycnodusH. pictetiHaqelA pycnodontid pycnodont.
HastichthysH. gracilisHaqel & NammouraA dercetid aulopiform.
HayenchelysH. germanusHjoulaAn eel.
HayolperichthysH. pectospinusHaqelA gladiopycnodontid pycnodont.
HemisauridaH. hakelensisHaqelA halecid aulopiform.
HensodonH. spinosusHaqelA coccodontid pycnodont
HeterothrissaH. signeuxaeHaqelA ctenothrissiform acanthomorph.
HgulichthysH. spinusHjoulaA trachichthyiform.
HumilichthysH. orientalisHjoulaA ctenothrissiform acanthomorph.
IchthyocerosI. spinosusHaqelA coccodontid pyncodont.
IchthyotringaI. delicataHaqel & HjoulaAn ichthyotringid aulopiform.
JoinvillichthysJ. kriwetiHaqel & HjoulaA gladiopycnodontid pycnodont.
J. lindstroemiHaqel
LebonichthysL. lewisiHjoulaA bonefish.
L. namourensisNammoura
LebrunichthysL. nammourensisNammouraA pachyrhizodontid.
LibanoberyxL. spinosusA trachichthyiform.
LissoberyxL. arambourgiHjoulaA trachichthyiform.
L. dayi
L. denticulatus
LuenchelysL. minimusNammouraAn eel.
MaraldichthysM. verticalisHaqelA gebrayelichthyid pycnodont.
MicrocaprosM. libanicusHjoulaA quaesitoberycid beryciform.
MonocerichthysM. scheuchzeriHaqel & HjoulaA gladiopycnodontid pycnodont.
NematonotusN. bottaeHaqel & NammouraA flagfin.
N. longispinusHjoula
'Nursallia''N.' goedeliHaqel, Hjoula & NammouraA pycnodontid pycnodont.
'N.' tethysensisNammoura and Hjoula
OmosomaO. sp./gen. nov. sp. nov.NammouraPossibly an indeterminate beardfish, potentially a new genus.
OrnategulumO. sardinioidesHaqel & NammouraA clupeomorph.
PalaeopantodonP. vandersypeniHaqelA pantodontid.
PankowskichthysP. libanicusHaqel & NammouraA gladiopycnodontid pycnodont.
PankowskipiscisP. haqelensisHaqelA pantodontid.
ParacentrusP. lebanonensisNammouraA holocentriform
P. sp. nov.
Paraclupeidae indet.nov. gen. 1NammouraA paraclupeid clupeomorph; two different genera.
nov. gen. 2.Hjoula
ParacoccodusP. woodwardiHaqelA coccodontid pyncodont.
PateropercaP. libanicaHjoulaA ctenothrissiform acanthomorph.
P. robusta
PattersonichthysP. delicatusHjoulaA ctenothrissiform acanthomorph.
PattersonoberyxP. pharsusHaqelA potential holocentriform.
PetalopteryxP. syriacusHaqelAn aphanepygid.
PetersichthysP. libanicusHaqelA pantodontid.
PharmacichthysP. numismalisHaqelA pharmacichthyid lamprimorph.
P. venenifer
PhoenicolepisP. arcuatusHjoulaA ctenothrissiform acanthomorph.
PhylactocephalusP. microlepisHaqel & HjoulaA halecid aulopiform.
PlectocretacicusP. claraeHaqelA potential tetraodontiform.
PlesioberyxP. discoidesHaqelAn indeterminate acanthopterygian.
P. maximus
PrognathoglossumP. kalassyiHaqelA pantodontid.
PrionolepisP. cataphractusHaqel & HjoulaA prionolepid aulopiform.
ProtobramaP. avusHaqel & HjoulaA protobramid tselfatiiform.
P. woodwardiHaqel
Protobramidae indet.nov. gen. nov. sp.NammouraA protobramid tselfatiiform.
PseudoberyxP. bottaeHaqel & HjoulaAn indeterminate teleost.
P. grandisHaqel
P. syriacusHaqel & Hjoula
PycnosteroidesP. levispinusHjoulaA pycnosteroidid lamprimorph.
QuaesitoberyxQ. minutusHjoulaA quaesitoberycid beryciform.
RhamphoichthysR. taxidiotisHaqelA billfish-like plethodid tselfatiiform.
RhinopycnodusR. gabriellaeHaqelA pycnodontid pycnodont.
RhynchodercetisR. hakelensisHaqel & HjoulaA dercetid aulopiform.
RostropycnodusR. gayetiHaqelA gladiopycnodontid pycnodont.
SardinioidesS. attenuatusHaqel & HjoulaA sardinioidid myctophiform.
S. minimus
S. pontivagus
SaurorhamphusS. giorgiaeNammouraAn enchodontid aulopiform.
ScombroclupeaS. diminutaNammouraA clupeomorph.
S. macrophthalmaHaqel & Hjoula
ScutatoclupeaS. bacchiaiNammouraA paraclupeid clupeomorph.
SedenhorstiaS. dayiHjoulaA tarpon.
S. libanicaHaqel
S. orientalisHjoula
SerrilepisS. minorNammouraA halecid aulopiform.
S. prymnostrigos
SigmapycnodusS. giganteusHaqelA pycnodontid pycnodont.
SorbinichthysS. elusivoNammouraA sorbinichthyid clupeomorph.
SpathiurusS. dorsalisHaqelAn ionoscopiform.
SpinascutichthysS. pankowskiaeNammouraAn enchodontoid aulopiform.
StanhopeichthysS. libanicusHjoulaA pachyrhizodontid.
StenoprotomeS. hamataHaqelA gladiopycnodontid pycnodont.
StichocentrusS. elegansHjoulaA holocentriform.
S. liratus
S. spinulosus
S. sp.
StichopteryxS. lewisiHaqel & HjoulaA trachichthyiform.
TelepholisT. tenuisHaqel & HjoulaA cheirothricid.
TrewavasiaT. carinataHaqelA coccodontid pycnodont.
TricerichthysT. wenziHaqelA gladiopycnodontid pycnodont.
TriplomystusT. nooraeNammouraA paraclupeid clupeomorph.
T. oligoscutatus
VolcichthysV. sp.HaqelAn indeterminate aulopiform or sardinioid myctophiform.
YpsiloichthysY. sibelleaeHaqelA teleost of uncertain affinities.

Lobe-finned fish

Sarcopterygians reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
MacropomoidesM. orientalisHaqel & HjoulaA latimeriid coelacanth.

Amniotes

Amniotes reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
AphanizocnemusA. libanensisAlthough the type locality is unknown, it is said to "almost certainly" originate from the Sannine Formation.A marine lizard.
EnantiophoenixE. electrophylaNammouraAn enantiornithine bird.
EupodophisE. descouensiNammouraAn early marine simoliophiid snake with hind limbs.
MicrotubanM. altivolansHjoulaAn azdarchoid pterosaur.
MimodactylusM. libanensisHjoulaAn istiodactyliform pterosaur.
PontosaurusP. kornhuberiNammouraA marine dolichosaurid lizard.
RhinochelysR. nammourensisNammouraA marine protostegid turtle. Adult and multiple hatchling remains known, suggesting a nesting site was nearby.
Theropoda indet.NammouraThe partial arm of a theropod dinosaur, the only non-avialan dinosaur material known from the formation. Now lost.

Invertebrates

Arthropods

Arthropods reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
AcanthochiranaA. smithwoodwardiHaqel & HjoulaAn aegerid prawn.
AcanthophoenicidesA. peterpaniA slipper lobster.
AegerA. libanensisHaqel & HjoulaAn aegerid prawn.
ArmacrangonA. edwardsiA glyphocrangonid shrimp.
?BentheogennemaB. phoeniciaA benthesicymid crustacean.
CallianassaC. sp.HjoulaA callianassid crustacean.
CalliaxinaC. hadjoulaensisHjoulaA callianassid crustacean.
CarinacarisC. antennaeA penaeid prawn.
C. teruzziHaqel & Hjoula
CarpopenaeusC. callirostrisHaqel & HjoulaA carpopenaeid prawn.
C. septemspinatusHaqel & Hjoula
CharbelicarisC. maronitesHjoulaA slipper lobster.
ChresmodaC. libanicaNammouraA water strider-like chresmodid insect.
CorazzatocarcinusC. hadjoulaeHjoulaA necrocarcinid crab.
CretaxiopsisC. libanoticaAn axiid crustacean.
CretacalliaxC. levantinaHjoulaA callianopsid ghost shrimp.
EpipenaeusE. abisaadorumAn carpopenaeid shrimp.
ErymaE. oscariHjoulaAn erymid "lobster".
GladicrangonG. coriaceaA glyphocrangonid shrimp.
GlobulacarisG. garassinoiHaqelA protozoeid thylacocephalan.
GlypheaG. damesiHjoulaA glypheoid crustacean.
HakelocarisH. vavassoriiHaqel & HjoulaA penaeid prawn.
HomolopsisH. aff. edwardsiiHjoulaA carrier crab.
JasusJ. sp.HjoulaA spiny lobster.
JoinvilleicarisJ. longirostrisA penaeid prawn.
LibanoaxiusL. beataeAn axiid crustacean.
LibanocarisL. curvirostraA penaeid prawn.
L. rogeriHaqel & Hjoula
LebanoaktassiaL. curiosaHjoulaAn aktassiid dragonfly.
LibanoaeshnaL. mikhaeliHaqelA libanoaeshnid dragonfly.
LibanobelostomaL. calineaeHaqelA giant water bug.
LibanocorduliaL. debieiHjoulaA libanocorduliid dragonfly
LibanogomphusL. lionelcaviniHjoulaA libanogomphid dragonfly.
LibanoliupanshaniaL. mimiHjoulaA liupanshaniid dragonfly
LinuparusL. secretanaeA spiny lobster.
MacropenaeusM. incertusHaqel & HjoulaA penaeid prawn.
MagilaM. cretacicaAn axiid crustacean.
MicrochelaM. rostrataHaqel & HjoulaA penaeid prawn.
MieroblattinaM. pacisNammouraA mesoblattinid cockroach.
NotahomarusN. hakelensisHaqel & HjoulaA true lobster.
OdontochelionO. bicarinatumA crangonid shrimp.
O. cretaceumHaqel & Hjoula
PalaeobenthonectesP. arambourgiA benthesicymid crustacean.
PalaeopalinurellusP. jbeilensisHaqelA slipper lobster.
PalaeopandalusP. komaiiA pandalid shrimp.
PalibacusP. praecursorHaqel & HjoulaA slipper lobster.
PalinurusP. teruzziiA spiny lobster.
ParacancrinosP. libanensisHjoulaA slipper lobster.
ParadollocarisP. vannieriHjoulaA dollocaridid thylacocephalan.
ParanecrocarcinusP. cf. milbourneiHjoulaA necrocarcinid crab. Specimen consumed by a catshark.
PhalangiopsisP. antoineiA chirostylid squat lobster.
P. rogeri
PhoeniceP. pasiniiHaqelA boxer shrimp.
PseudastacusP. dubertretiHaqel & HjoulaA stenochirid "lobster".
PustulinaP. cretaceaHaqel & HjoulaAn erymid "lobster".
StriadiogenesS. frigerioiHjoulaA diogenid hermit crab.
TachypleusT. syriacusHjoulaA horseshoe crab.
TelamonocarcinusT. gambulatusHaqel & HjoulaA dorippid crab.
ThylacocarisT. schramiHjoulaA dollocaridid thylacocephalan.

Molluscs

Molluscs reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
AllocriocerasA. cf. annulatumAn anisoceratid ammonite.
BoreopeltisB. smithiHaqel or HjoulaA plesioteuthid coleoid.
DorateuthisD. syriacaHaqel & HjoulaA plesioteuthid coleoid.
EdmunditeuthisE. bacchiaiHaqel & HjoulaA trachyteuthid coleoid.
GlyphidopsisG. waageiHaqelA trachyteuthid coleoid.
GlyphiteuthisG. abisaadorumHaqel & HjoulaA trachyteuthid coleoid.
G. freijiiHaqel, Hjoula & Nammoura
G. libanoticaHaqel & Hjoula
KeuppiaK. levanteHjoulaAn early octopus. 2 or 3 species known from the same locality
K. hyperbolaris
K. sp.
MantellicerasM. mantelliHaqel"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"
RachiteuthisR. acutaliHjoulaA palaeololiginid coleoid.
R. donovaniHâqel
StyletoctopusS. annaeHâqelAn early octopus.
TeudopsiniaT. haasiHaqelA teudopsid coleoid.
SyrionautilusS. libanoticusHjoulaA cymatoceratid nautiloid.

Echinoderms

Echinoderms reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
AntedonA. pinnulataHaqelA crinoid.
Aspidochirotida indet.HjoulaA sea cucumber.
GeocomaG. libanoticaHaqelA brittle star.

Annelids

Annelids reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
DidoneD. pulcherrimaA didonid bristleworm.
EunicitesE. diopatroidesA eunicid bristleworm.
E. falcatus
E. joinvillei
E. mariacristinae
FerragutiaF. cenomanianaA bristleworm.
LumbriconereitesL. garassinoiA bristleworm.
L. hadjulae
PalaeoaphroditeP. libanicaAn aphroditid bristleworm.
PhoeniciarabellaP. caesarisA lumbrinerid bristleworm.
P. orensanzi
P. pinnulata
TeruzziaT. gryphoeidesA lumbrinerid bristleworm.
T. pezzolii
T. pusilla
T. sagittifera
RollinschaetaR. myoplenaHaqel & HjoulaA fireworm.

Plants

Plants reported from the Sannine Formation
GenusSpeciesPresenceNotesImages
Aryskumiacf. A. zelkovifoliaNammouraDicot known from a poorly preserved leaf with ovate blade.
NammouriaN. gracilisAn aquatic fern of uncertain affinities.
NammourophyllumN. altingioidesA dicot known from leaves, a possible member of the Altingiaceae.
NupharanthusN. cretaceaA dicot known by a preserved flower, possibly a member of the Nympheales.
ParvileguminophyllumP. sp.Dicot known from a complete leaflet, possibly a member of the Fabaceae.
'Phragmites'P. sp.A monocot of uncertain affinities, vaguely similar to the modern reed.
Platycaryeae gen. et sp. indet.Infructescence of bracteate fruits resembling Platycarya americana.
PseudolarixP. sp.A conifer related to the modern golden larch.
PseudotorelliaP. sp.A gymnosperm of uncertain affinities.
PterisP. sp.A pteridacean fern.
SapindopsisS. anhouryiA platanacean, related to the modern plane trees.
S. libanensis

See also