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''Odontochelys semitestacea'' (meaning "toothed turtle with a half-shell") is a Late
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest per ...
relative of
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
s. Before ''
Pappochelys ''Pappochelys'' ( meaning "grandfather turtle" in Greek) is an extinct genus of diapsid reptile possibly related to turtles. The genus contains only one species, ''Pappochelys rosinae'', from the Middle Triassic of Germany, which was named by pa ...
'' was discovered and '' Eunotosaurus'' was redescribed, ''Odontochelys'' was considered the oldest undisputed member of Pantestudines (i.e. a
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
-turtle). It is the only known species in the genus ''Odontochelys'' and the family Odontochelyidae.


Discovery

''Odontochelys semitestacea'' was first described from three 220-million-year-old specimens excavated in
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest per ...
deposits in
Guizhou Guizhou (; Postal romanization, formerly Kweichow) is a landlocked Provinces of China, province in the Southwest China, southwest region of the China, People's Republic of China. Its capital and largest city is Guiyang, in the center of the pr ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
. The locale of its discovery at one time was the Nanpanjiang Trough basin, a shallow marine environment surrounded on three sides by land. These deposits preserve an ecosystem known as the
Guanling biota The Xiaowa Formation is a Carnian-age geological formation found in southern China. It is a sequence of limestone and marls from the Carnian stage of the Triassic. Its lower section was previously known as the Wayao Formation or Wayao Member of ...
, which was dominated by marine reptiles.


Description

''Odontochelys'' differed grossly from modern turtles. Modern turtles have a horny
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for fo ...
without teeth in their mouth. In contrast, ''Odontochelys'' fossils were found to have had teeth embedded in their upper and lower jaws. One of the most striking features of turtles, both modern and prehistoric alike, are their dorsal shells, forming an armored carapace over the body of the animal. ''Odontochelys'' only possessed the bottom portion of a turtle's armor, the plastron. It did not yet have a solid
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
as most other turtles do. Instead of a solid carapace, ''Odontochelys'' possessed broadened ribs like those of modern turtle embryos that still have not started developing the
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
plates of a carapace. It reached nearly in total body length. Aside from the presence of teeth and the absence of a solid carapace, a few other skeletal traits distinguish ''Odontochelys'' as basal compared to other turtles,
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
and otherwise. The point of articulation between the dorsal ribs and the vertebrae is decidedly different in ''Odontochelys'' than in later turtles. In a comparison of
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
proportions, the skull of ''Odontochelys'' is far more elongated pre-orbitally (in front of the eyes) compared to other turtles. The tail of the prehistoric turtle was longer in proportion to its body than other turtles. In addition, the
transverse process 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 ...
es found in the tail are not fused such as in later turtles. Also, the
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eith ...
e of the examined specimens were identified to lack acromion processes. Taken together, these anatomical differences have been interpreted by the discoverers to mean that ''Odontochelys'' has some of the most primitive features ever seen in a turtle and is somewhat of a
transitional fossil 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 ...
. Evidence that the plastron evolved before the carapace, as indicated by the lack of carapace in ''Odontochelys semitestacea'', is often viewed as an indication of the aquatic origin of turtles. The fossil was found in marine deposits, further supporting that the primitive turtle frequented shallow marine water. Since it is generally accepted that the shell arose to provide protection against predators, the semi-aquatic nature of turtles and the development of the plastron complement each other. Ancestral turtles with protection on their underside are more protected from predators that attack from below. Based on this interpretation, the development of the carapace was likely driven in a land animal. Reisz and Head (2008), however, have a different interpretation on the same specimen. Instead, they suggest that the carapace on ''O. semitestacea'' was in fact present; it just lacked ossification of some of its dermal components. With this interpretation, the authors suggest that either turtle shells originally evolved in aquatic environments, or this fossil represents the earliest turtle transferring from terrestrial environments to marine habitats.


Paleoecology

Even though the ''Odontochelys'' specimens were found in marine deposits, there is uncertainty over whether it was a primarily aquatic reptile, in transition from aquatic to terrestrial habitats, or fully terrestrial. Li ''et al.'' (2008) argued that ''Odontochelys'' lived in coastal or freshwater environments based on its hand proportions, which were similar to those of modern turtle species which lived in small, slow-moving bodies of water. Joyce (2015) supported a terrestrial origin for turtles based on his study of ''Eunotosaurus''. His study noted that the hands of ''Odontochelys'' were not comparable to any modern turtles due to the retention of a
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...
("primitive") feature: four
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. ...
(finger bones) in the third and fourth fingers, rather than three. While the total relative finger lengths of ''Odontochelys'' do line up with those of semiaquatic freshwater turtles, the individual phalanges (finger bones) themselves were short and stout, much more similar to those of modern
tortoise Tortoises () are reptiles of the family Testudinidae of the order Testudines (Latin: ''tortoise''). Like other turtles, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like oth ...
s. Aquatic turtles achieved long hands by lengthening their phalanges, while ''Odontochelys'' retained long hands due to its plesiomorphic phalangeal count, similar to that of other reptiles (including terrestrial ones). In addition, Joyce (2015) argued that even if ''Odontochelys'' was semiaquatic, it probably would have avoided open marine waters due to a lack of efficient adaptations for swimming. On the other hand, a
pathological Pathology is the study of the causes and effects of disease or injury. The word ''pathology'' also refers to the study of disease in general, incorporating a wide range of biology research fields and medical practices. However, when used in th ...
study of ''Odontochelys'' performed by Rothschild & Naples (2015) discovered that both the left and right humeri (forearm bones) of the
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...
specimen (IVPP 13240) of ''Odontochelys'' had been degraded near the shoulder sockets. The study rejected explanations such as
weathering Weathering is the deterioration of rocks, soils and minerals as well as wood and artificial materials through contact with water, atmospheric gases, and biological organisms. Weathering occurs '' in situ'' (on site, with little or no movement ...
or a wound-induced bone infection, arguing that it would not have made sense for the shoulder area to degrade before the rest of the forelimbs, since the shoulder was more well-protected during life and after death. Instead, the study argued that
decompression sickness Decompression sickness (abbreviated DCS; also called divers' disease, the bends, aerobullosis, and caisson disease) is a medical condition caused by dissolved gases emerging from solution as bubbles inside the body tissues during decompressio ...
was responsible for the injury. This condition has been observed in diving animals which are forced to make rapid ascents within a deep marine environment. Similar injuries have been reported in other fossilized marine reptiles, and their presence in ''Odontochelys'' supports the idea that it lived in an open marine environment. Modern sea turtles utilize behavioral tactics to avoid rapid ascension within water, which may also indicate that ''Odontochelys'' had not yet acquired the same behaviors to defend against decompression sickness.


References


External links


Figures and Tables
from the above-referenced article ''An ancestral turtle from the Late Triassic of southwestern China'' in the journal ''
Nature Nature, in the broadest sense, is the physical world or universe. "Nature" can refer to the phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. Although humans are ...
''
Oldest Turtle Found; May Crack Shell-Evolution MysteryFirst known turtle swam on the half shell
{{Taxonbar, from=Q131325 Pantestudines Prehistoric reptile genera Triassic reptiles of Asia Prehistoric animals of China Triassic China Fossils of China Fossil taxa described in 2008 Transitional fossils