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''Polycotylus'' is a
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
plesiosaur The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared ...
within the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Polycotylidae Polycotylidae is a family of plesiosaurs from the Cretaceous, a sister group to Leptocleididae. Polycotylids first appeared during the Albian stage of the Early Cretaceous, before becoming abundant and widespread during the early Late Cretaceous. ...
. The
type species In 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 species that contains the biological type specimen ...
is ''P. latippinis'' and was named by American paleontologist
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 1869. Eleven other species have been identified. The name means 'much-cupped vertebrae', referring to the shape of the vertebrae. It lived in 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, ...
of North America toward the end of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
. One fossil preserves an adult with a single large fetus inside of it, indicating that ''Polycotylus'' gave live birth, an unusual adaptation among reptiles.


History

Edward Drinker Cope named ''Polycotylus'' from the
Niobrara Formation The Niobrara Formation , also called the Niobrara Chalk, is a geologic formation in North America that was deposited between 87 and 82 million years ago during the Coniacian, Santonian, and Campanian stages of the Late Cretaceous. It is compose ...
in
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 ...
in 1869. 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 several ...
bones from which he based his description were fragmentary, representing only a small portion of the skeleton. A more complete skeleton was later found in Kansas and was described in 1906. A nearly complete skeleton was found in 1949 from 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 ...
in
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 ...
, but was not described until 2002. A new species, ''P. sopozkoi'', from Russia was described in 2016.


Description

Like all plesiosaurs, ''Polycotylus'' was a large marine reptile with a short tail, large flippers, and a broad body. It has a short neck and a long head, and was about long and weighed . It has more neck vertebrae than other polycotylids, however. ''Polycotylus'' is thought to be a basal polycotylid because it has more vertebrae in its neck (a feature that links it with long-necked ancestors) and its
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
has a more primitive shape. The long
ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from Naples. It is the largest of the Phlegrean Islands. Roughly trapezoidal in shape, it measures approximately east to west ...
of the pelvis are a distinguishing feature of ''Polycotylus'', as are thick teeth with striations on their surfaces, a narrow pterygoid bone on the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
and a low
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
on top of the skull.


Classification

Unlike some better-known long-necked plesiosaurs like ''
Plesiosaurus ''Plesiosaurus'' (Greek: ' ('), near to + ' ('), lizard) is a genus of extinct, large marine sauropterygian reptile that lived during the Early Jurassic. It is known by nearly complete skeletons from the Lias Group, Lias of England. It is disting ...
'' and ''
Elasmosaurus ''Elasmosaurus'' (;) is a genus of plesiosaur that lived in North America during the Campanian stage of the Late Cretaceous period, about 80.5million years ago. The first specimen was discovered in 1867 near Fort Wallace, Kansas, US, and was se ...
'', ''Polycotylus'' had a short neck. This led to it being classified as a
pliosaur Pliosauroidea is an extinct clade of plesiosaurs, known from the earliest Jurassic to early Late Cretaceous. They are best known for the subclade Thalassophonea, which contained crocodile-like short-necked forms with large heads and massive toot ...
, a marine reptile within the superfamily Pliosauroidea, closely related to true plesiosaurs (which belong to the superfamily Plesiosauroidea). ''Polycotylus'' and other polycotylids superficially resemble pliosaurs like ''
Liopleurodon ''Liopleurodon'' (; meaning 'smooth-sided teeth') is an extinct genus of large, carnivorous marine reptile belonging to the Thalassophonea, a clade of short-necked pliosaurid plesiosaurs. ''Liopleurodon'' lived from the Callovian Stage of the Mi ...
'' and ''
Peloneustes ''Peloneustes'' (meaning "mud swimmer") is a genus of pliosaurid plesiosaur from the Middle Jurassic of England. Its remains are known from the Peterborough Member of the Oxford Clay Formation, which is Callovian in age. It was originally descri ...
'' because they have short necks, large heads, and other proportions that differ from true plesiosaurs. As
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses became common in the last few decades, the classification of ''Polycotylus'' and other plesiosaurs have been revised. In 1997, it and other polycotylids were reassigned as close relatives of long-necked elasmosaurids. In a 2001 study, ''Polycotylus'' was classified as a
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
cryptocleidoid plesiosaur closely related to
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
plesiosaurs like ''
Cryptocleidus ''Cryptoclidus'' ( ) is a genus of plesiosaur reptile from the Middle Jurassic period of England, France, and Cuba. Discovery ''Cryptoclidus'' was a plesiosaur whose specimens include adult and juvenile skeletons, and remains which have bee ...
''. Below is a cladogram from a 2004 study which supported a similar classification: In 2007, ''Polycotylus'' was placed in a new subfamily of polycotylids called Polycotylinae. Another newly described polycotylid called '' Eopolycotylus'' from
Glen Canyon Glen Canyon is a natural canyon carved by a length of the Colorado River, mostly in southeastern and south-central Utah, in the United States. Glen Canyon starts where Narrow Canyon ends, at the confluence of the Colorado River and the Dirty D ...
,
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
, was found to be the closest relative of ''Polycotylus''. Below is a cladogram from the 2007 study:


Paleobiology


Reproduction

A fossil of ''P. latippinis'' catalogued LACM 129639 includes an adult individual with a single
fetus A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal deve ...
inside it. LACM 129639 was found in Kansas during the 1980s and was in storage at the
Natural History Museum of Los Angeles County The Natural History Museum of Los Angeles County is the largest natural and historical museum in the western United States. Its collections include nearly 35 million specimens and artifacts and cover 4.5 billion years of history. This large coll ...
until it was described in 2011. The length of the fetus is around , which is about 32 percent of the length of the mother. Gestation was probably two thirds complete based on what is known of the fetal development of related
nothosaur Nothosaurs (order Nothosauroidea) were Triassic marine sauropterygian reptiles that may have lived like seals of today, catching food in water but coming ashore on rocks and beaches. They averaged about in length, with a long body and tail.F. v. ...
s. This fossil suggests that ''Polycotylus'' was viviparous, giving live birth (as opposed to laying eggs).
Viviparity Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
, or live birth, may have been the most common form of reproduction in plesiosaurs, as they would have had difficulty laying eggs on land. Their bodies are not adapted to movement on land, and paleontologists have long hypothesized that they must have given birth in water. Other marine reptiles such as
ichthyosaur Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...
s also gave live birth, but LACM 129639 was the first direct evidence of vivipary in plesiosaurs. The lives of ''Polycotylus'' and other plesiosaurs were
K-selected In ecology, ''r''/''K'' selection theory relates to the selection of combinations of traits in an organism that trade off between quantity and quality of offspring. The focus on either an increased quantity of offspring at the expense of indivi ...
, meaning that few offspring were born to each individual but those that were born were cared for as they mature. Because it gave birth to a single large offspring, the mother ''Polycotylus'' probably gave it some form of parental care for it to survive. F. Robin O'Keefe, one of the describers of LACM 129639, suggested that the social lives of plesiosaurs were "more similar to those of modern dolphins than other reptiles." K-selected life-history strategies are also seen in mammals and some lizards, but are unusual among reptiles. Examination of the fetus of ''Polycotylus'' indicates that while in the womb, plesiosaurs sacrificed fetal bone strength for accelerated growth rates. Histological analysis and comparisons with another plesiosaur, ''
Dolichorhynchops ''Dolichorhynchops'' is an extinct genus of polycotylid plesiosaur from the Late Cretaceous (early Turonian to late Campanian stage) of North America, containing three species, ''D. osborni'', ''D. bonneri'' and ''D. tropicensis'', as well as a ...
'', showed that some plesiosaur infants were up to forty percent the length of the mother when born, and that infant plesiosaurs may have had some compromised swimming abilities.


See also

*
List of plesiosaur genera This list of plesiosaurs is a comprehensive listing of all genera that have ever been included in the order Plesiosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considered inv ...
*
Timeline of plesiosaur research This timeline of plesiosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, taxonomic revisions, and cultural portrayals of plesiosaurs, an order of marine reptiles that flourished duri ...


References

{{Taxonbar, from=Q1886121 Late Cretaceous plesiosaurs of North America Polycotylids Fossil taxa described in 1869 Taxa named by Edward Drinker Cope Sauropterygian genera