Protostega
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''Protostega'' ('first roof') 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
sea turtle Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerh ...
containing a single species, ''Protostega gigas''. Its fossil remains have been found in the Smoky Hill Chalk formation of western
Kansas Kansas ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the west. Kansas is named a ...
(''Hesperornis'' zone, dated to 83.5 million years agoCarpenter, K. (2003). "Vertebrate Biostratigraphy of the Smoky Hill Chalk (Niobrara Formation) and the Sharon Springs Member (Pierre Shale)." ''High-Resolution Approaches in Stratigraphic Paleontology'', 21: 421-437. ), time-equivalent beds of the Mooreville Chalk Formation of
Alabama Alabama ( ) is a U.S. state, state in the Southeastern United States, Southeastern and Deep South, Deep Southern regions of the United States. It borders Tennessee to the north, Georgia (U.S. state), Georgia to the east, Florida and the Gu ...
and
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
beds of the Rybushka Formation (
Saratov Oblast Saratov Oblast is a federal subjects of Russia, federal subject of Russia (an oblast), located in the Volga Federal District. Its administrative center is the types of inhabited localities in Russia, city of Saratov. As of the 2021 Russian cens ...
,
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
). Fossil specimens of this species were first collected in 1871, and named by
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontology, paleontologist, comparative anatomy, comparative anatomist, herpetology, herpetologist, and ichthyology, ichthyologist. Born to a wealthy Quaker fam ...
in 1872. With a total length of , it is the second-largest sea turtle that ever lived, second only to the giant ''
Archelon ''Archelon'' is an extinct marine turtle from the Late Cretaceous, and is the largest turtle ever to have been documented, with the biggest specimen measuring from head to tail and in body mass. It is known only from the Pierre Shale and ...
'', and one of the three largest turtles of all time alongside ''
Archelon ''Archelon'' is an extinct marine turtle from the Late Cretaceous, and is the largest turtle ever to have been documented, with the biggest specimen measuring from head to tail and in body mass. It is known only from the Pierre Shale and ...
'' and '' Gigantatypus''.


Discovery and history

The first known ''Protostega'' specimen (YPM 1408) was collected on July 4 by the 1871 Yale College Scientific Expedition, close to Fort Wallace and about 5 months before Cope arrived in Kansas. However, the fossil that they found was never described or named. It was not named until 1872, when E. D. Cope found and collected the first identified specimen of ''Protostega gigas'' in the Kansas chalk in 1871. A variety of bones were found in yellow Cretaceous chalk from a bluff near Butte Creek.


Paleoenvironment

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 ...
was marked by high temperatures, with large epicontinental seaways. During the Mid-to-Late Cretaceous period the
Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, or the Western Interior Sea) was a large inland sea (geology), inland sea that existed roughly over the present-day Great Plains of ...
covered the majority of North America and would connect to the Boreal and Tethyan oceans at times. Within these regions are where the fossil of ''Protostega gigas'' have been found.


Description

''Protostega'' is known to have reached up to in length. A specimen from the upper Taylor Marl is even larger, at in carapace length and in total length. Despite lacking its head and three limbs, it is well-preserved. Cope's ''Protostega gigas'' discovery revealed that their shell had a reduction of
ossification 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 ...
that helped these huge animals with streamlining in the water and weight reduction. The
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 ...
was greatly reduced and the disk only extending less than halfway towards the distal ends of the ribs. Cope described other greatly modified bones in his specimen including an extremely long coracoid process that reached all the way to the pelvis and a humerus that resembled a '' Dermochelys,'' creating better movement of their limbs. Edward Cope described ''Protostega gigas'' as having a large jugal that reached to the quadrate along with a thickened pterygoid that reached to the mandibular articulating surface of the quadrate. The fossil featured a reduction in the posterior portion of the vomer where the palatines meet medially. Another fossilized specimen showed that a bony extension, that would have been viewed as a beak, was lacking in the ''Protostega'' genus. The premaxillary beak was much shorter than that of ''
Archelon ''Archelon'' is an extinct marine turtle from the Late Cretaceous, and is the largest turtle ever to have been documented, with the biggest specimen measuring from head to tail and in body mass. It is known only from the Pierre Shale and ...
''. In front of the orbital region the skull was elongated with a broadly-roofed temporal region. The jaws of the fossil showed a large crushing surface. The quadrato-jugal was triangular with a posterior edge that was concave, with the entire bone being convex from distal view. The squamosal appeared to have a concave formation on the surface at the upper end of the quadrate. In Cope's fossil the mandible was preserved almost perfectly and from this he recorded that the jaw was very similar to the
Cheloniidae Cheloniidae is a family of typically large marine turtles that are characterised by their common traits such as, having a flat streamlined wide and rounded shell and almost paddle-like flippers for their forelimbs. They are the only sea turtles ...
and the dentary had a broad for above downward with a concave surface, marked by deep pits in the dentary. Cope concluded that these animals were most likely
omnivore An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...
s and consumed a diet of hard shelled crustacean creatures, due to the long symphysis of its lower jaw. ''Protostega'' also likely fed on seaweed and jellyfish or scavenged on floating carcasses as well, like modern turtles.


Classification

The classification of ''Protostega'' was complicated at best. The specimen that Cope discovered in Kansas was hard to evaluate with the preservation condition. The fossil shared many characteristics with the genus '' Dermochelys'' and the family Cheloniidae''.'' Cope wrote about the characteristics that distinctly separated this particular species from the two controversial groups. The differences he described were that the fossil had a reduced or lacking amount of dermal ossification on the back, the articulation of the pterygoid and quadrates, the presence of a presplenial bone in the jaw, a lack of an articular process on the back side of the nuchal, simple formation of the radial process on the humerus, and a peculiar bent formation of the xiphiplastra. He concluded that ''Protostega gigas'' was an intermediate form between ''Dermochelys'' and Cheloniidae.


Paleobiology

Examining the bone tissue microstructure (osteohistology) of ''Protostega'' revealed growth patterns similar to modern leatherback sea turtles with rapid growth to large body size. Leatherbacks lack a typical reptile metabolism, instead having high resting metabolic rates and the ability to hold a body temperature higher than their surroundings. If ''Protostega'' had similar bone growth patterns to leatherbacks, it is hypothesized that they both had a similar metabolism. This rapid growth to adult body size in sea turtles would also indicate rapid growth to reproductive maturity, which would have been a great advantage in their survival. However, comparing ''Protostega'' to its more basal relative '' Desmatochelys'' shows that not all protostegids had the same growth patterns. This indicates that rapid growth to large size evolved late within the lineage, perhaps in response to the evolution of large
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
s like ''
Tylosaurus ''Tylosaurus'' (; "knob lizard") is a genus of Russellosaurina, russellosaurine mosasaur (an extinct group of predatory marine Squamata, lizards) that lived about 92 to 66 million years ago during the Turonian to Maastrichtian stages of the Late ...
''. Given uncertainties in the phylogenetic placement of protostegids relative to living sea turtles, it is unclear if the evolution of rapid growth rates and possible elevated metabolism were convergent with modern leatherbacks or if the two were more closely related.


See also

* List of Turtles *
Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, or the Western Interior Sea) was a large inland sea (geology), inland sea that existed roughly over the present-day Great Plains of ...
*
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 ...


References

{{Taxonbar, from=Q1051251 Protostegidae Late Cretaceous turtles of North America Prehistoric turtle genera Taxa named by Edward Drinker Cope Mooreville Chalk Extinct turtles Fossil taxa described in 1872