Protostega Gigas
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''Protostega'' ('first roof') 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 extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
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, loggerhead, ...
containing a single species, ''Protostega gigas''. Its fossil remains have been found in the
Smoky Hill Chalk The Smoky Hill Chalk Member of the Niobrara Chalk formation is a Cretaceous conservation Lagerstätte, or fossil rich geological formation, known primarily for its exceptionally well-preserved marine reptiles. Named for the Smoky Hill River, the S ...
formation of western
Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the ...
(''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 The Mooreville Chalk is a geological formation in North America, within the U.S. states of Alabama and Mississippi, which were part of the subcontinent of Appalachia. The strata date back to the early Santonian to the early Campanian stage of t ...
of
Alabama (We dare defend our rights) , anthem = "Alabama (state song), Alabama" , image_map = Alabama in United States.svg , seat = Montgomery, Alabama, Montgomery , LargestCity = Huntsville, Alabama, Huntsville , LargestCounty = Baldwin County, Al ...
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. Campanian s ...
beds of the
Rybushka Formation The Rybushka Formation is a Campanian geologic Formation (geology), formation in the Penza Oblast, Penza and Saratov Oblasts of European Russia. Pterosaur, fish and invertebrate fossils have been recovered from the formation. Fossil content The ...
(
Saratov Oblast Saratov Oblast (russian: Сара́товская о́бласть, ''Saratovskaya 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 ...
,
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
). 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, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interested ...
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 Dakota Pierre Shal ...
'', and one of the three largest turtle of all time along ''
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 Dakota Pierre Shal ...
'' 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 wasn't until
E. D. Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy intereste ...
found and collected the first 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 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'', the ...
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, and the Western Interior Sea) was a large inland sea that split the continent of North America into two landmasses. The ancient sea, ...
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 reveled 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 Dorsum (biology), 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 tor ...
was greatly reduced and the disk only extending less than half way 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 The leatherback sea turtle (''Dermochelys coriacea''), sometimes called the lute turtle or leathery turtle or simply the luth, is the largest of all living turtles and the heaviest non-crocodilian reptile, reaching lengths of up to and weights ...
''. Creating better movement of their limbs. Skull Structure: Edward Cope described the uniqued ''Protostega gigas'' to have 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 a bony extension, that would have been viewed as a beak, was lacking in the ''Protostega'' genus. The premaxillary beak was very 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 Dakota Pierre Shal ...
''. In front of the orbital region was elongated with broadly roofed temporal region. The jaws of the fossil showed a large crushing -surfaces. The quadrato-jugal was triangular with a posterior edge that was concave and the entire bone was 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 ''Chelonidae'' 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 has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
s and consumed a diet of hard shelled crustacean creatures, due to the long symphysis of its lower jaw. Along with probably consisted of 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 two other recorded genus named ''
Dermochelys The leatherback sea turtle (''Dermochelys coriacea''), sometimes called the lute turtle or leathery turtle or simply the luth, is the largest of all living turtles and the heaviest non-crocodilian reptile, reaching lengths of up to and weights ...
'' and '' Chelonidae.'' 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, presplenial bone in the jaw was present, no 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 genus ''Protostega'' and species ''Protostega gigas'' was an intermediate form of the two groups ''Dermochelys'' and Chelonidae.


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, and the Western Interior Sea) was a large inland sea that split the continent of North America into two landmasses. The ancient sea, ...
*
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'', the ...


References

{{Testudines Protostegidae Late Cretaceous turtles of North America Prehistoric turtle genera Taxa named by Edward Drinker Cope Mooreville Chalk Extinct turtles