Drazinderetes
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''Drazinderetes'' is a large bodied genus of soft shell turtle from the
Middle Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "da ...
Drazinda Formation of
Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's fifth-most populous country, with a population of almost 243 million people, and has the world's second-lar ...
. Its presence in the shallow marine deposits of the Drazinda Formation suggests that ''Drazinderetes'' may have been a partially or fully marine animal. Indetermined trionychine remains from the same formation may suggest that ''Drazinderetes'' could have been among the largest known turtles, with one entoplastron indicating a potential length of 1.5 to 2.1 meters. ''Drazinderetes'' currently consists of only a single species: ''Drazinderetes tethyensis''.


Discovery and naming

The first surveys of the Drazinda Formation were conducted in 1993 and 1996 by the
Geological Survey of Pakistan Geological Survey of Pakistan (GSP) is an independent executive scientific agency to explore the natural resources of Pakistan. Main tasks GSP perform are Geological, Geophysical and Geo-chemical Mapping of Pakistan. Target of these mapping are res ...
and the
University of Michigan , mottoeng = "Arts, Knowledge, Truth" , former_names = Catholepistemiad, or University of Michigania (1817–1821) , budget = $10.3 billion (2021) , endowment = $17 billion (2021)As o ...
. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
(GSP-UM 3 195) of ''Drazinderetes'', a nearly complete carapace, was found during the 1996 exploration of the area west of Satta Post in sediments of
Bartonian The Bartonian is, in the ICS's geologic time scale, a stage or age in the middle Eocene Epoch or Series. The Bartonian Age spans the time between . It is preceded by the Lutetian and is followed by the Priabonian Age. Stratigraphic defini ...
age (39 - 38 Ma) that also yielded
archaeocetes Archaeoceti ("ancient whales"), or Zeuglodontes in older literature, is a paraphyletic group of primitive cetaceans that lived from the Early Eocene to the late Oligocene (). Representing the earliest cetacean radiation, they include the initial ...
and primitive sea cows. Several other remains assigned to softshell turtles have been found from the same sediments, however they largely don't overlap with the type specimen and can thus not be directly referred to it. This includes GSP-UM 3019, an exceptionally large entoplastron, and GSP-UM 3 185, a small hypoplastron possibly belonging to a juvenile specimen. GSP-UM 3 185, a carapace fragment, is the only specimen to overlap with the holotype. It is slightly larger and heavily weathered, with smooth edges that do not match what is observed in the type specimen. Given the insufficient material, these specimen could only be designated as Trionychidae indet. and like the holotype, all these specimen have been collected in 1996. The name derives from the Drazinda Formation and "eretes", Greek for rower (mirroring the name of ''
Aspideretes ''Nilssonia'' is a genus of softshell turtles (family Trionychidae) from rivers, streams, ponds, and lakes in South Asia and Burma. In many treatments, it is monotypic, with the single species Burmese peacock softshell (''N. formosa''). However, ...
''). The species name derives from the Tethys Sea which the animal likely inhabited.


Description

The shell of ''Drazinderetes'' shows a fully preserved nuchal plate followed by 8 pairs of costal plates. It's roughly oval in shape when viewed from above, with the right half notably better preserved in the holotype specimen. The shell expands sideways through the 2nd costal and reaches its greatest width (700 mm) from the 3rd to 5th costal plates. After the 5th costal the carapace tapers until the final plate of the shell. The back end of the shell shows a distinct concave margin. On top of the shell, between the nuchal and first costal plate sits a hexagonal, raised bony plate that corresponds in position and in its connection to the first
thoracic vertebra In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae and they are intermediate in size between the cervical ...
to the preneural plate in other softshell turtles. The following neural plates however are largely obscured due to the poor preservation of the carapace's midline. The entire carapace is notably sculpted with a surface covered by tightly intertwined patterns that are in turn overlain by concentric rings that become more irregular towards the center of the shell. The inside of the shell shows a single pair of costiform processes that are fused to the nuchal plate. The processes do not protrude beyond the shell's edge and overlap with the first pair of thoracic
ribs The rib cage, as an enclosure that comprises the ribs, vertebral column and sternum in the thorax of most vertebrates, protects vital organs such as the heart, lungs and great vessels. The sternum, together known as the thoracic cage, is a semi- ...
. The ribs themselves possess thin heads and do not extend very far from the edge of the costal plates unlike in some other softshell turtles. They are generally rectangular or convex in outline.


Size

Unlike in many other testudines, the shell of trionychid turtles is composed only partly of the bony carapace, with cartilage making up 20% (
Apalone ''Apalone'' is a genus of turtles in the family Trionychidae. Species of ''Apalone'' are native to North America. Geographic range Most ''Apalone'' species are restricted to the United States, though the range of the spiny softshell, ''A. spini ...
) to 45% (
Malayan softshell turtle The Malayan softshell turtle (''Dogania subplana'') is a species of softshell turtle in the family Trionychidae. It is monotypic in its genus. Geographic range It is found in Brunei, Indonesia, Java, Kalimantan, Malaysia, Myanmar, the Philippin ...
) of the entire carapace length depending on the species. Since the exact relationship between ''Drazinderetes'' (and GSP-UM 3019) and other softshell turtles are largely unknown, Head and colleagues instead opted to calculate both the minimum and maximum extend of cartilage as seen in modern taxa. The holotype specimen preserves a bony carapace that measures mm in length. In accordance with modern species, the cartilage may add up to a total carapace length between and . For GSP-UM 3019, an exceptionally large specimen that may belong to the same species, a total carapace length of 1.2 meters was extrapolated based on the known plastron remains. Following the same method applied to the ''Drazinderetes'' holotype, this would yield a total length between to meters. This not only places GSP-UM 3019 as the largest recorded trionychid, outsizing '' Axestemys'', an Eocene softshell turtle from the
Bridger Formation The Bridger Formation is a geologic formation in southwestern Wyoming. It preserves fossils dating back to the Ypresian Epoch of the Paleogene Period. The formation was named by American geologist Ferdinand Vandeveer Hayden for Fort Bridger, whi ...
of
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to the s ...
(bony carapace length of ), but as one of the largest known turtles. Even the lower estimates rival modern leatherback sea turtles, while upper estimates are comparable to large protostegids such as Archelon. Merely the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
podocnemidid ''
Stupendemys ''Stupendemys'' is an extinct genus of freshwater side-necked turtle, belonging to the family Podocnemididae. It is the largest freshwater turtle known to have existed, with a carapace over 2 meters long. Its fossils have been found in norther ...
'' is larger. Ignoring this specimen, ''Drazinderetes'' is still notably larger than any extant softshell turtle.


Phylogeny

Upon its description, ''Drazinderetes'' was assigned to the Trionychinae due to the reduced costiform processes typical for the clade as well as characters of the carapace. Analysis conducted by Head ''et al.'' showed that ''Drazinderetes'' shared characteristics with the Indo-Asian trionychine lineage and place it as a close relative of the dubious genus ''
Aspideretes ''Nilssonia'' is a genus of softshell turtles (family Trionychidae) from rivers, streams, ponds, and lakes in South Asia and Burma. In many treatments, it is monotypic, with the single species Burmese peacock softshell (''N. formosa''). However, ...
''. In 2017, Georgalis and Joyce recovered the following phylogentic tree in their research on old world softshell turtles.


Paleoenvironment

Given the lack of evidence that would suggest that the bones of ''Drazinderetes'' had been washed into the marine sediments of the Drazinda Formation
postmortem An autopsy (post-mortem examination, obduction, necropsy, or autopsia cadaverum) is a surgical procedure that consists of a thorough examination of a corpse by dissection to determine the cause, mode, and manner of death or to evaluate any d ...
, it is deemed likely by Head and colleagues that ''Drazinderetes'' was instead a fulltime or at least occasional inhabitant of saltwater environments. Typically freshwater inhabitants wandering into the sea is not unheard of in modern softshell turtles, notably the giant softshell turtle. The unusually large size suggested by some of the specimen may support this fully marine lifestyle, as large body size is often found in marine turtles (tho not exclusive to such a lifestyle). Additionally, if all specimen belong to different ontogenetic stages of a single species, then the presence of younger and fully grown animals in a single environment would furthermore speak in favor of this hypothesis. This would make ''Drazinderetes'' the first fully marine softshell turtle.


See also

*
Largest prehistoric animals The largest prehistoric animals include both vertebrate and invertebrate species. Many of them are described below, along with their typical range of size (for the general dates of extinction, see the link to each). Many species mentioned might ...
*
List of largest reptiles The list of largest reptiles includes living crocodilians that reach a length of and more, such as Orinoco crocodile (''Crocodylus intermedius''), American crocodile (''C. acutus''), Nile crocodile (''C. niloticus''), saltwater crocodile (''C. ...


References

{{Testudines Extinct turtles Prehistoric turtle genera Trionychinae Prehistoric turtles of Asia Eocene turtles Fossils of Pakistan Fossil taxa described in 1999 Cenozoic reptiles of Asia