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''Eupodophis'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
snake Snakes are elongated limbless reptiles of the suborder Serpentes (). Cladistically squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales much like other members of the group. Many species of snakes have s ...
from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
period. It has two small hind legs and is considered a transitional form between Cretaceous
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
s and limbless snakes. The feature, described as vestigial, was most likely useless to ''Eupodophis''. The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
''Eupodophis descouensi'' was named in 2000 and resides now in the paleontology section of the Mim Museum 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 International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
-age
limestone Limestone is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...
near the al-Nammoura village in
Lebanon Lebanon, officially the Republic of Lebanon, is a country in the Levant region of West Asia. Situated at the crossroads of the Mediterranean Basin and the Arabian Peninsula, it is bordered by Syria to the north and east, Israel to the south ...
belonging to the
Sannine Formation 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 wit ...
.


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 east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
Tethys Ocean The Tethys Ocean ( ; ), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean during much of the Mesozoic Era and early-mid Cenozoic Era. It was the predecessor to the modern Indian Ocean, the Mediterranean Sea, and the Eurasia ...
. It had a laterally compressed body and short, paddle-like tail. The
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e and
rib In vertebrate anatomy, ribs () are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the thoracic cavity, enabling the lungs to expand and thus facilitate breathing by expanding the ...
s of ''Eupodophis'' are pachyostotic, 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 (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
s and
phalanges The phalanges (: phalanx ) are digit (anatomy), digital bones in the hands and foot, feet of most vertebrates. In primates, the Thumb, thumbs and Hallux, big toes have two phalanges while the other Digit (anatomy), digits have three phalanges. ...
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 strength of the magnetic field which bends the particle beam i ...
x-ray An X-ray (also known in many languages as Röntgen radiation) is a form of high-energy electromagnetic radiation with a wavelength shorter than those of ultraviolet rays and longer than those of gamma rays. Roughly, X-rays have a wavelength ran ...
s at the
European Synchrotron Radiation Facility The European Synchrotron (ESRF) is a joint research facility situated in Grenoble, France, supported by 19 countries (13 member countries: Belgium, Denmark, Finland, France, Germany, Italy, the Netherlands, Norway, Russia, Spain, Sweden, Switz ...
in
Grenoble Grenoble ( ; ; or ; or ) is the Prefectures in France, prefecture and List of communes in France with over 20,000 inhabitants, largest city of the Isère Departments of France, department in the Auvergne-Rhône-Alpes Regions of France, region ...
,
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
. The researchers determined that the hind limb on one skeleton was 0.8 inches long, with an "unmistakable"
fibula The fibula (: fibulae or fibulas) 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. ...
,
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
and
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
. 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 Mexico, Mexican state of Sonora. It is a heavy, slow-moving reptile, up to long, and it is the only ve ...
,
green iguana The green iguana (''Iguana iguana''), also known as the American iguana or the common green iguana, is a large, arboreal, mostly Herbivory, herbivorous species of lizard of the genus ''Iguana''. Usually, this animal is simply called the 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 West African Nile monitor, one species is also found in south America as an invasive species. A ...
. 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 outer and inner parts of the bone are compact), the limb bones remain light. This lightness is also seen in the bones of terrestrial lizards, suggesting that the limbs had not been part of the overall adaptation of the skeleton for an aquatic lifestyle.


Paleobiology

The loss of limbs in ''Eupodophis'' may have been the result of changes in
Hox gene Hox genes, a subset of homeobox, homeobox genes, are a gene cluster, group of related genes that Evolutionary developmental biology, specify regions of the body plan of an embryo along the craniocaudal axis, head-tail axis of animals. Hox protein ...
s, genes that specify the development specific regions of the body. Because Hox genes are involved in determining specific features of the axial skeleton, the loss of limbs would also result in the loss of cervical (tail) vertebrae that are near them. This loss is seen in ''Eupodophis'' and modern snakes but not legless lizards, which may be far less common because some other factor besides Hox genes were involved in the loss of their limbs. The loss of digits on the hind limbs may be explained by a low number of cells in the
limb bud The limb bud is a structure formed early in vertebrate limb development. As a result of interactions between the ectoderm and underlying mesoderm, formation occurs roughly around the fourth week of development. In human embryonic development, the ...
during embryonic development. The loss of forelimbs and reduction of hind limbs in ''Eupodophis'' was likely an adaptation for swimming. While living snakes usually employ undulatory movement for moving over land, sinuous movements are also an effective means of moving through water. Large, well-developed limbs increase drag on swimming animals, so the limbs of ''Eupodophis'' and other early snakes may have become vestigial to save energy and make movement more efficient. No vestigial limbs whatsoever are present in the modern species, which lack transitional species remnants. What few species have them protrude as tiny spurs. Most herpetologists agree snakes diverged from lizard forms by discarding impediments to effective sheltering in subterrestrial environments. Most modern snakes live above the surface.


See also

*
List of transitional fossils 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 ...
Other known fossil snakes with legs: *'' Pachyrhachis'' *'' Haasiophis'' *'' Najash''


References


External links


The Cenomanian: stage of hindlimbed snakes

Ancient serpent shows its leg
Jonathan Amos, BBC News {{Taxonbar, from=Q1954854 Cretaceous reptiles of Asia Cretaceous snakes Transitional fossils