''Eupodophis'' is an

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...

snake Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more ...

from the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...

period. It has two small hind legs and is considered a

transitional form A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross a ...

between Cretaceous

lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia altho ...

s and limbless snakes. The feature, described as vestigial, was most likely useless to ''Eupodophis''. The

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...

''Eupodophis descouensi'' was named in 2000 and resides now in the paleontology section of the

Mim Museum The Mim Museum is a private museum in Beirut, Lebanon. The museum displays more than 2000 minerals, representing 450 different species from 70 countries, and is considered one of the most significant private collections of minerals in the world. I ...

in Beirut, Lebanon. The specific name is dedicated to the French naturalist Didier Descouens. The fossilized specimen from which the description of the type species was based was 85 cm (33.5 in) long and is approximately 92 million years old. It was found in

Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...

-age

limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...

near the al-Nammoura village in

Lebanon Lebanon ( , ar, لُبْنَان, translit=lubnān, ), officially the Republic of Lebanon () or the Lebanese Republic, is a country in Western Asia. It is located between Syria to the north and east and Israel to the south, while Cyprus li ...

belonging to the

Sannine Formation The Sannine Formation, also called the Sannine Limestone, is a Cretaceous geologic formation in Lebanon. Description It is primarily Cenomanian in age. The formation laterally varies from east to west; the western lowland "coastal" sequence i ...

Description

''Eupodophis'' was a marine snake that lived in the

Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...

Tethys Ocean The Tethys Ocean ( el, Τηθύς ''Tēthús''), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean that covered most of the Earth during much of the Mesozoic Era and early Cenozoic Era, located between the ancient continents ...

. It had a laterally compressed body and short, paddle-like tail. The

vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...

e and

rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...

s of ''Eupodophis'' are

pachyostotic Pachyostosis is a non-pathological condition in vertebrate animals in which the bones experience a thickening, generally caused by extra layers of lamellar bone. It often occurs together with bone densification (osteosclerosis), reducing inner ca ...

, or thickened, as an adaptation to a marine lifestyle. The pelvic bones are small and weakly attached to each other. Tarsal bones are present but reduced in size and form. The

metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...

s and

phalanges The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...

of the foot are absent. The fossil skeleton of ''Eupodophis'' was analyzed using

synchrotron A synchrotron is a particular type of cyclic particle accelerator, descended from the cyclotron, in which the accelerating particle beam travels around a fixed closed-loop path. The magnetic field which bends the particle beam into its closed p ...

x-ray An X-ray, or, much less commonly, X-radiation, is a penetrating form of high-energy electromagnetic radiation. Most X-rays have a wavelength ranging from 10 picometers to 10 nanometers, corresponding to frequencies in the range 30&nb ...

s at the

European Synchrotron Radiation Facility The European Synchrotron Radiation Facility (ESRF) is a joint research facility situated in Grenoble, France, supported by 22 countries (13 member countries: France, Germany, Italy, the UK, Spain, Switzerland, Belgium, the Netherlands, Denmark, ...

Grenoble lat, Gratianopolis , commune status = Prefecture and commune , image = Panorama grenoble.png , image size = , caption = From upper left: Panorama of the city, Grenoble’s cable cars, place Saint- ...

France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...

. The researchers determined that the hind limb on one skeleton was 0.8 inches long, with an "unmistakable"

fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity is ...

tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...

and

femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...

. One limb was visible on the surface of the fossil while the other was hidden within the limestone. The scans were compared with similar ones taken of the limbs of extant lizards including the

Gila monster The Gila monster (''Heloderma suspectum'', ) is a species of venomous lizard native to the Southwestern United States and the northwestern Mexican state of Sonora. It is a heavy, typically slow-moving reptile, up to long, and it is the only v ...

, green iguana, and several species of

monitor lizard Monitor lizards are lizards in the genus ''Varanus,'' the only extant genus in the family Varanidae. They are native to Africa, Asia, and Oceania, and one species is also found in the Americas as an invasive species. About 80 species are recogn ...

While they are very small in comparison to limbed reptiles, the hind limbs of ''Eupodophis'' possess much of the same anatomy as modern lizards. This suggests that the bones of ''Eupodophis'' became reduced in size through a change in the rate of bone growth, not major anatomical changes. The lack of thickening at either end of the limb bones suggests that growth had stopped occurring in the limbs at one point in the animal's lifetime. While the vertebrae and ribs of ''Eupodophis'' are pachyostotic and osteosclerotic (meaning that the

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