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 during the Mesozoic Era. The first scientifically documented plesiosaur fossils were discovered during the early 19th century by Mary Anning.[1] Plesiosaurs were actually discovered and described before dinosaurs.[2] They were also among the first animals to be featured in artistic reconstructions of the ancient world, and therefore among the earliest prehistoric creatures to attract the attention of the lay public.[3] Plesiosaurs were originally thought to be a kind of primitive transitional form between marine life and terrestrial reptiles. However, now plesiosaurs are recognized as highly derived marine reptiles descended from terrestrial ancestors.[4]
Early researchers thought that plesiosaurs laid eggs like most reptiles. They commonly imagined plesiosaurs crawling up beaches and burying eggs like turtles. However, later opinion shifted towards the idea that plesiosaurs gave live birth and never went on dry land: the biggest genera may have been too heavy to go on land at all, but smaller genera could've been capable.[5] Plesiosaur locomotion has been a source of continuous controversy among paleontologists.[6] The earliest speculations on the subject during the 19th century saw plesiosaur swimming as analogous to the paddling of modern sea turtles. During the 1920s opinion shifted to the idea that plesiosaurs swam with a rowing motion.[7] However, a paper published in 1975 that once more found support for sea turtle-like swimming in plesiosaurs.[8] This conclusion reignited the controversy regarding plesiosaur locomotion through the late 20th century.[9] In 2011, F. Robin O'Keefe and Luis M. Chiappe concluded the debate on plesiosaur reproduction, reporting the discovery of a gravid female plesiosaur with a single large embryo preserved inside her.[10]
Parkinson coined the name Plesiosaurus priscus for some of the remains used by de la Beche and Conybeare as the basis for Plesiosaurus. This species is currently regarded as of dubious taxonomic value.[14]
Mary Anning discovered a nearly complete Plesiosaurus skeleton near Lyme Regis. This specimen would later be catalogued as BMNH 22656.[15]
c. December
Around the same time as the discovery of BMNH 22656, another Plesiosaurus specimen was discovered at the same site. The specimen was donated to the Oxford University museum and is probably the specimen known today as OXFUM J.10304.[16]
Conybeare described the new species name Plesiosaurus dolichodeirus for the Plesiosaurus discovered by Anning. As the first species name given to a distinctive and well preserved Plesiosaurus skeleton it has come to be regarded as both the type specimen of Plesiosaurus dolichodeirus specifically and of the genus Plesiosaurus as a whole.[15][17]
De la Beche illustrated a work titled "Duria Antiquior", meaning "A More Ancient Dorset" for fossil hunter Mary Anning. This work, which prominently features plesiosaurs, has been regarded as the first attempt to accurately reconstruct the Mesozoic world through an artistic medium.[3]
Cast of the flipper of Pliosaurus brachydeirus. The species was named by Owen in 1841.Skeleton of the "Barrow Kipper" Atychodracon megacephalus specimen
The trustees of the British Museum of Natural History bought the type specimen of Plesiosaurus from the estate of the first duke of Buckingham, Richard Glenville. The museum catalogued the specimen as BMNH 22656.[16]
An army surgeon named Dr. Theophilus Turner discovered the fossils of a large animal in the Pierre Shale of Kansas, USA. The remains represented the first nearly complete plesiosaur specimen from North America. Turner gave some of its vertebrae to a member of the Union Pacific Railroad's survey named John LeConte. LeConte sent the vertebrae to Edward Drinker Cope for study. Cope recognized the find as a significant plesiosaur discovery and wrote to Turner asking him to excavate and ship the fossils to him.[24]
March, mid
An 1869 life restoration of Elasmosaurus (with the head still on the wrong end) confronting the theropod dinosaur Laelaps (now Dryptosaurus) platyurus.
Cope erected the new genus and species Elasmosaurus platyurus for the fossils sent by Turner in a rushed descriptive manuscript written within two weeks of obtaining them.[25]
March 24
Cope presented his findings regarding Elasmosaurus to a meeting of the Academy of Natural Sciences in Philadelphia, Pennsylvania.[26]
Cope prepared a preprint for the Transactions of the American Philosophical Society of his Elasmosaurus description, including reconstruction of the animal with a short neck and very long tail. The manuscript was then distributed to other scholars.[26]
Cope's mentor Joseph Leidy gave a presentation reporting his recent discovery that Cope's reconstruction of Elasmosaurus positioned the skull at the end of the tail rather than the end of the neck.[26]
Leidy's discovery embarrassed Cope, who began spreading notice of an unspecified error in his Elasmosaurus description with an offer to replace it with a corrected version and its second volume.[28]
November
O. C. Marsh collected a better an additional specimen of Polycotylus in Kansas that was better preserved than the type described by Cope. The specimen is now catalogued as YPM 1125.[29]
Skull of Cryptoclidus. C. eurymerus was originally described by Phillips in 1871.
Cope imagined elasmosaurs feeding by craning their necks above the water and striking downward at fish long distances from their bodies.[30]
B. F. Mudge discovered ten articulated vertebrae in the Fairport Chalk of Kansas that he mistook for ichthyosaur remains. These fossils are now catalogued as KUVP 1325.[31]
B. F. Mudge discovered fragments of a large elasmosaur skeleton in the Fort Hays Limestone of Kansas.[32]
Mudge and Williston excavated the remains another large Kansan plesiosaur, this one from the Smoky Hill Chalk.[32] The specimen may be a Styxosaurus snowii and is currently catalogued as YPM 1644. It was the first plesiosaur Mudge had ever found with gastroliths and the first plesiosaur encountered by Williston in general.[33]
Hector reported the presence of elasmosaur remains in New Zealand.[34]
Seeley published a paper intended to help improve the state of science's understanding of plesiosaur shoulder girdle anatomy, which had been muddled by the poor preservation of the fossils many early paleontologists had to rely on for their observations.[35]
Mudge discussed the gastroliths of YPM 1644 in a scientific publication.[33] He concluded that plesiosaur exploited gastroliths to assist in breaking down food the way many modern birds and reptiles do.[37]
Oxford acquired the Misses Philpot collection, which included the type specimen of Plesiosaurus macromus. The museum catalogued this specimen as OXFUM J.28587.[38]
The British Museum of Natural History purchased the Edgerton collection, which included the complete Plesiosaurus dolichodeirus jaw now known as BMNH R.255.[18]
The Smithsonian bought a partial plesiosaur skeleton from Charles Sternberg. The specimen is now catalogued as USNM 4989 and would later serve as the type specimen of the new genus and species Brachauchenius lucasi.[39]
In an article published in the New York Herald, Marsh brought up Cope's anatomic reversal of Elasmosaurus.[28]
E. P. West excavated a skull and partial neck belonging to the elasmosaur that would come to be named Styxosaurus snowii. The specimen is now catalogued as KUVP 1301.[42]
Williston described the species now known as Styxosaurus snowii.[19]
Marsh described the new species Pantosaurus striatus.[21]
Charles H. Sternberg obtained two large elasmosaur vertebrae that would later serve as the type specimen of Elasmosaurus sternbergi. The specimen is now catalogued as KUVP 1312.[43]
F. W. Cragin discovered a partial plesiosaur skeleton and associated gastroliths in what is now recognized as the Kiowa Shale. This specimen is now catalogued as KUVP 1305 and would later be named Plesiosaurus mudgei.[44]
Williston argued that plesiosaurs ingested gastroliths only accidentally or to relieve "food craving[s]".[37] However, he also observed that the rocks used as gastroliths were more similar to rocks 400–500 miles away in Iowa or the Black Hills of South Dakota than those of the local geology.[45]
With guidance from Edward Drinker Cope, paleo-artist Charles R. Knight illustrated an Elasmosaurus platyurus eating a fish. The elasmosaur's neck was erroneously looped into an anatomically impossible figure 8 configuration that evoked the image of "a python grasping at its prey".[46]
George F. Sternberg discovered the plesiosaur specimen now known as KUVP 1300 that would later serve as the type specimen of Dolichorhynchops osborni.[48]
Williston made several changes to plesiosaur taxonomy. One of these was the description of the new genus and species Brachauchenius lucasi, whose type specimen was a partial skeleton discovered in Kansas. This specimen is now catalogued as USNM 4989.[39] He also described the new species Trinacromerum anonymum based on the vertebral series discovered by Mudge in 1872. This specimen is now known as KUVP 1325.[31] Lastly, Williston regarded Plesiosaurus mudgei as a junior synonym of the species Plesiosaurus gouldii.[44] He also commented on the ongoing debate regarding plesiosaur gastroliths, acknowledging the possibility that they were used for ballast while also maintaining openness to his 1893 suggestion that the stones were ingested accidentally.[49]
Barnum Brown hypothesized that plesiosaurs used their gastroliths in a gizzard-like organ to grind up their invertebrate prey since they had no grinding or crushing teeth to do that job for them.[49]
Harvard paleontologist Charles R. Eastman, "offended" by Brown's claim that plesiosaurs had a gizzard, criticized the idea in print.[49]
Williston responded to Eastman, reasserting the evidence for plesiosaur gastroliths by noting that by this time at least 30 specimens containing them had been found.[50]
Williston described several new taxa and specimens. One of these was the new species Elasmosaurus nobilis.[32] Williston also described the new Elasmosaurus species E. sternbergi based on the vertebrae discovered by Charles H. Sternberg in 1893. He remarked that these fossils were the largest elasmosaur vertebrae that he had ever seen.[43] Lastly, Williston described Marsh's Polycotylus, YPM 1125.[29]
Williston observed that the semicircular canals inside a plesiosaur's ear were well developed, giving them a good sense of balance and coordination.[53]
The Smithsonian obtained the Tylosaurus specimen with preserved polycotylid stomach contents from Charles Sternberg.[54] The Tylosaurus is catalogued as USNM 8898 and its last supper as USNM 9468.[55]
Sternberg observed that being contained in the stomach of a mosasaur might have helped ensure the preservation of the polycotylid now known as USNM 9468 by protecting it from scavenging sharks.[55]
Harvard University dispatched a fossil hunting expedition to Queensland, Australia. In Army Downs they discovered a nearly complete specimen of Kronosaurus.[57]
The University of Nebraska State Museum bought the elasmosaur specimen discovered by Sternberg and Walker in 1931. The specimen is now catalogued as UNSM 1195.[56]
A specimen of Trinacromerum was discovered in a roadside exposure of the Greenhorn Formation in Kansas. The specimen is now catalogued as KUVP 5070.[31]
A large pliosaur skeleton was found on the banks of Russia's Volga River. However, the specimen was damaged during the excavation and only the skull and chest region were successfully extracted.[59]
A complete specimen of Plesiosaurus conybeari, including preserved soft tissues, was destroyed in a bombing raid against Bristol. Fortunately, a cast of the specimen survived in the British Museum.[60]
White described the new species Seeleyosaurus holzmadensis.[61]
Alfred Sherwood Romer helped mount the Kronosaurus discovered in Queensland by the 1930s Harvard expedition for the University's Museum of Comparative Zoology. The poorly preserved bones required a significant amount of plaster for the restoration, earning the specimen the mocking nickname "Plasterosaurus".[57] The final mount was 42 feet long, probably due to Romer overestimating the number of vertebrae in its spine; a more likely length is about 35 feet.[63]
Fossil hunters Robert and Frank Jennrich serendipitously discovered a partial Brachauchenius skeleton when looking for sharks teeth.[52]
Shuler, like Williston in 1914, found elasmosaurs to have relatively inflexible necks.[36] He also found elasmosaurs to have stereoscopic vision, which would have been useful for hunting small prey.[53]
October
George Sternberg excavated the Brachauchenius discovered by the Jennriches. This specimen, now known as FHSM VP-321, was both larger and better preserved than the Brachauchenius type specimen. Although it was put on display soon after discovery, it would not be described for the scientific literature for nearly 50 years.[52]
Barney H. Newman and Lambert Beverly Halstead Tarlo argued that long-necked plesiosaur flippers could only move horizontally, and while maneuverable, they were confined to surface waters by an inability to dive.[65]
South Australianopal miners John and Molly Addyman discovered a plesiosaur whose remains had been converted to opal.[66]
Paul Johnston discovered plesiosaur fossils in a roadside exposure of the Greenhorn Formation in Kansas.[67] During the excavation the dig site was scouted by two suspicious men. After a break from digging the Johnston team returned to find all of the fossils crudely extracted from the rock except for a flipper that the team had reburied. Based on the flipper, the stolen plesiosaur could be identified as Trinacromerum bentonianum.[68]
Robinson publishes follow up research to her previous publication on plesiosaur locomotion.[9] This second paper notably concluded that plesiosaurs were incapable of leaving the water.[69]
Michael Alan Taylor published a paper concluding that plesiosaurs would have been capable of moving on land after all because their spinal column was too arched for their lungs to collapse.[70]
Samuel F. Tarsitano and Jürgen Riess published a paper harshly critical of Robinson's previous work on plesiosaur locomotion. However, while criticizing Robinson's work they were reluctant to make any positive claims of their own, concluding that the details of plesiosaur locomotion were "unknown".[9]
Richard A. Thulborn published the results of his recent re-examination of the purported plesiosaur embryos discovered by Harry Govier Seeley. Thulborn concluded that Seeley's supposed embryos were actually nodules of mudstone and shale derived from sediments that once filled in a crustaceanburrow system and were not even animal body fossils.[40]
Delair described the new species Bathyspondylus swindoniensis.[21]
A South Australian opal miner named Joe Vida discovered the skeleton of a juvenile plesiosaur whose remains had converted to opal. Its preparator, Paul Willis nicknamed it Eric. An entrepreneur named Sid Londish bought the specimen and funded its preparation, but went bankrupt. When the specimen was put up for auction fear spread that a potential buyer might break the specimen down for its gemstone value. A television drive was arranged on behalf of the Australian Museum. The Museum succeeded in raising 340,000 dollars to buy the specimen, whose gemstone value was about $300,000. Eric was later identified as a specimen of Leptocleidus.[22]
Liopleurodon teeth share traits with those of killer whales (skeleton pictured)
Wiffen and Moisley described the new species Tuarangisaurus keysei.[21]
Judith Massare published an analysis of plesiosaur feeding habits. She concluded that the long-necked plesiosauroids ate soft prey. Liopleurodon and its relatives, on the other hand, had teeth resembling those of killer whales and probably ate larger, bonier prey.[51]
Orville Bonner discovered a specimen of Dolichorhynchops osborni that was later seen to preserve developing young inside it.[71]
Judy Massare analyzed Mesozoic marine reptile swimming abilities and found that long-necked plesiosaurs would have been significantly slower than pliosaurs due to excess drag incurred from the length of the neck.[72]
The Los Angeles County Museum of Natural History acquired the Dolichorhynchops osborni specimen discovered by Bonner and catalogued it as LACMNH 129639.[44]
Beverly Halstead published a paper suggesting that plesiosaurs swam using all four flippers paired with an undulatory motion of the body comparable to a sea lion's.[73]
Nakaya reported the presence of elasmosaur remains in Japan.[34]
Stewart noted a relative paucity of plesiosaur fossils from the lower portions of the Smoky Hill Chalk in a manuscript for the Society of Vertebrate Paleontology.[75]
J. D. Stewart, accompanied by Everhart, discovered a nearly complete Dolichorhynchops rear flipper in the lower Smoky Hill Chalk. Unfortunately it was too late to correct the erroneous statements in his aforementioned paper regarding the supposed rarity of plesiosaurs in the lower Smoky Hill Chalk. The flipper is now catalogued as LACMNH 148920.[77]
Location of Lawrence in the U.S. state of Kansas
October
Stewart's paper, complete with his now-erroneous statements, was published in the Niobrara Chalk Excursion Guidebook in honor of the society's 50th anniversary meeting in Lawrence that year.[75]
Ralph E. Molnar published suspicion that the "Kronosaurus queenslandicus" specimen discovered by the Harvard expedition might be a distinct species.[47]
Several Elasmosaurus vertebrae and gastroliths were found near the site where the type specimen of the genus itself was excavated.[64]
Cruikshank and others hypothesized that plesiosaurs could smell and taste water that "passively" flowed through their nasal passages while they swam.[53]
Everhart discovered some fragmentary plesiosaur fossils in the lower Smoky Hill Chalk of Kansas. Some of the fossils seemed to have been partially digested. The remains were later catalogued as FHSM VP-13966.[75]
Everhart showed the partially digested fossils to J. D. Stewart, who recognized them as pieces of a plesiosaur skull. The fossils are now catalogued as [clarification needed][77]
Everhart and his wife helped excavated a Styxosaurus snowii specimen in Kansas. During the dig Mrs. Everhart discovered an additional partial plesiosaur skeleton.[78]
Robert T. Bakker argued that Dolichorhynchops and its relatives became the most common kind of short-necked plesiosaur in the Cretaceous Western Interior Seaway after the ichthyosaurs became extinct. Further, they convergently evolved many traits similar to those of ichthyosaurs, such as long snouts and large eyes.[80]
Tony Thulborn and Susan Turner examined the crushed skull of the long-necked plesiosaur Woolungasaurus and found the presence of tooth marks left by some giant predator. They hypothesized that a Kronosaurus was the culprit.[81]
Storrs, like Williston and Shuler before him, argued that long-necked plesiosaurs had relatively inflexible necks.[36]
Rothschild and Martin reported the presence of the remains of a fossilized fetus preserved in the abdomen of a Dolichorhynchops osborni.[82]
Glenn W. Storrs formally described the world's smallest plesiosaur for the scientific literature.[74]
South Dakota School of Mines paleontologist James Martin excavated and described a very young pliosaur specimen.[74] The specimen was in what would have been the middle of the Cretaceous Western Interior Seaway. It is more plausible to think that this young individual was born in the seaway itself rather than surviving the extremely long swim from shore. So, the specimen provided circumstantial evidence for live birth and possibly even parental care by pliosaurs in the Cretaceous Western Interior Seaway.[83] However, since there were no adult fossils present this interpretation is speculative. Further, the fossil site had been heavily vandalized before discovery by scientists. A large hole was found near the baby pliosaur that could have once held the bones of its mother or other pod members.[84]
An amateur fossil hunter named Simon Carpenter discovered a 7-foot-long Pliosaurus brachyspondylus skull in claypits of the Blue Circle cement works near Westbury, England. More of the skeleton was found in the vicinity and this specimen came to be regarded as the best preserved pliosaur ever found;[85] it is held by Bristol Museum.[86]
Ken Carpenter recognized the plesiosaur specimen discovered by Pamela Everhart in 1992 as one of the largest known specimens of Dolichorhynchops osborni, now catalogued as CMC VP-7055.[78]
Carpenter published a review of the Cretaceous short-necked plesiosaurs known from western North America. In this paper he both revised these plesiosaurs' taxonomy as well as offering observations on their biostratigraphy and evolution.[88] Carpenter described the new genus and species Plesiopleurodon wellesi.[21] He also argued against the prevailing trend to treat DolichorhynchopsTrinacromerum as taxonomic synonyms by observing that they could be distinguished based on their skull anatomy.[89] However, he did conclude that the Trinacromerum species T. anonymum and T. willistoni were junior synonyms of T. bentonianum.[31]
In his remarks on short-necked plesiosaur evolution, Carpenter argued that polycotylids were more closely related to long-necked plesiosaurs than pliosaurs.[90] He observed that Trinacromerum bentonianum seems to have existed from the late Cenomanian to the Turonian. This represents a span of time approximating 3.3 million years. He found Dolichorhynchops osborni to have had an even longer lifespan, from the middle Turonian to the early Campanian., or roughly 4 million years. His research also suggested that there was a span of time during the life of the Western Interior Seaway in which it was not inhabited by polycotylids.[29]
He also reported that the Dolichorhynchops specimen KUVP 40001 from the Pierre Shale of South Dakota may have achieved the extraordinary length of 23 feet.[34] The large size of the Pierre Shale Dolichorhynchops compared to those of the earlier Smoky Hill Chalk suggested to Carpenter that these plesiosaurs were evolving larger body sizes over time. In fact the Pierre Shale specimens of Dolichorhynchops were nearly as large as Brachauchenius lucasi.[78] Carpenter described a particularly large specimen of that latter taxon in this paper as well, specifically FHSM VP-321.[52] His study of Brachauchenius led him to concur with Williston that it was closely related to Liopleurodon ferox.[52]
Arthur R. I. Cruickshank, David M. Martill, and Leslie Noe formally named the new Oxford Clay pliosaur Pachycostasaurus dawni. The researchers noticed that its bones were very dense. So dense, they speculated it would naturally sink in the water and spent most of its time feeding on soft bodied animals living near the seafloor.[87]
Liggett and others reported the discovery of a giant plesiosaur flipper from the Greenhorn Limestone of Kansas. Although a significant portion of the specimen was missing, it implied a life length of more than 2 m. The researchers tentatively attributed the flipper to Brachauchenius lucasi. The specimen is now catalogued as FHSM VP-13997.[78]
Tamaki Sato and Kazushige Tanabe reported that a partial long necked plesiosaur skeleton had been discovered on the banks of Hokkaido's Obirashibe River. Although they could not identify its exact species, the specimen was significant because its stomach contents provided clues to long-necked plesiosaur diets. This individual's gut preserved the beaks of thirty small ammonites.[92]
John A. Long bemoaned the fact that the putative "Kronosaurus queenslandicus" uncovered by a Harvard team during the early 1930s had still not been formally described for the scientific literature.[47]
Michael Everhart and Glenn Storrs excavated additional Elasmosaurus ribs, vertebrae and gastroliths at the site of the 1991 discovery.[64]
Long reported the presence of elasmosaur remains in Australia.[34]
Carpenter published a summary of the elasmosaur fossils discovered in the Smoky Hill Chalk.[42]
Storrs published a revision of Elasmosaurus taxonomy.[93] He reinterpreted the Elasmosaurus nobilis type specimen as indeterminate elasmosaurid remains.[32] He also reinterpreted the "Elasmosaurus" sternbergi type specimen as two cervical and one dorsal vertebrae rather than two dorsal vertebrae as Williston had reported in his original description. However, Storrs did agree that it was an elasmosaur specimen rather than a pliosaur as argued by Welles in 1952.[43]
Lingham-Soliar argued that plesiosaur hind-flippers weren't mobile or muscular enough to help propel them through the water.[95]
Everhart published a study of the gastroliths associated with the elasmosaur specimen KUVP 129744 from Kansas. The specimen was associated with roughly 13.1 kg of gastroliths. The largest of these was 17 cm long and 1.4 kg in weight. Everhart would later compare its size to that of a softball and observe that not only was it one of the largest known plesiosaur gastroliths, but also one of the largest gastroliths from any animal.[37]
November
The Advertiser, a newspaper based in Adelaide, Australia bought the Addyman opalized plesiosaur specimen for $25,000 and donated it to the South Australian Museum. A paleontologist at the museum named Ben Kear identified it as a member of the genus Leptocleidus. The two foot long specimen was the smallest specimen of the genus ever found and probably a baby.[66]
David J. Cicimiurri and Michael J. Everhart published a study of the Styxosaurus snowii specimen NJSM 15435, which preserved both stomach contents and gastroliths.[37] Among the stomach contents were remains of the bony fish Enchodus.[53] By this point in time at least fifteen different plesiosaur specimens were known with preserved stomach contents.[96] The researchers observed that the Enchodus remains preserved in NJSM 15435 were an example of shifting dietary preferences in plesiosaurs, who fed primarily on cephalopods for most of their evolutionary history, before coming to rely more heavily on fishes during the Late Cretaceous.[53]
They also noted that some of NJSM 15435's gastroliths were scarred by rounded chips and arc-shaped marks. These were likely inflicted by contact with other gastroliths during the churning of the animal's stomach, and constituted physical evidence that plesiosaurs used their gastroliths to help break down their food during digestion.[97] Cicimurri and Everhart disputed the hypothesis that plesiosaurs used their gastroliths for ballast on the grounds that swallowing and vomiting such stones would be relatively difficult for the long-necked forms and their feeding grounds may have been hundreds of miles from sources of stones.[98]
Everhart resumed the study of the partially digested plesiosaur skull bones, FHSM VP-13966. He sought the expertise of Ken Carpenter due to his relevant 1996 paper on short-necked plesiosaurs. Carpenter identified the bones as probable Dolichorhynchops remains.[31]
Noe published another study of Pachycostasaurus. He changed his mind regarding its diet. Where previously he believed it to feed on soft-bodied animals, the robust and "heavily ornamented" build of its teeth suggested it fed on harder, bonier prey.[87]
Everhart was forced to cancel plans to examined the Tylosaurus specimen USNM 8898 and its polycotylid dinner USNM 9468 due to the September 11th terrorist attacks.[55]
November
Everhart was finally able to examine the tylosaur specimen with the polycotylid stomach contents.[55]
Robin O'Keefe publishes a series of papers examining the aspect ratios of plesiosaur flippers and comparing them to the wings of birds, bats, and aircraft. O'Keefe concluded that long-necked plesiosaurs were long-distance swimmers, while pliosaurs were more maneuverable.[100]
Michael Everhart examined FHSM VP-398 and found Sternberg's original note revealing that these fossils had been collected at the same site as the 1991 Elasmosaurus discovery. Everhart realized that the remains discovered there collectively represented most of the bones that had been missing from the Elasmosaurus type specimen. He inferred that they may represent fragments that fell off of the decomposing type carcass while it was adrift, before its final burial and fossilization.[64]
An elasmosaur specimen with over 600 associated gastroliths was discovered in the Pierre Shale of Nebraska. The specimen is now catalogued as UNSM 1111–002.[37]
December 30
The BBC began making sensational claims about the Aramberri pliosaur, claiming that the unidentified pliosaur was a Liopleurodon, that it was 65 feet long, that its teeth were the size of machetes, and its jaws powerful enough to crush granite.[101]
The University of Nebraska led the excavation of a long-necked plesiosaur skeleton on local Santee Sioux land.[102] The Santee people requested that the skeleton be mounted and displayed with a plaque acknowledging them as the source of the fossils and as having given permission for the museum to display the remains. However, the museum claims it could not honor the request as it did not have the funding to mount the skeleton for display, and it further claimed that the land the fossils were recovered from was of "disputed" ownership.[103]
Mulder and others reported the presence of elasmosaur remains in Europe.[34]
Sato described the new genus and species Terminonatator ponteixensis. In his study of the animal's skeleton, he found that the vertebral discs in the neck were flat on both sides and packed tightly together. He estimated that there would have been only about 0.5 cm of cartilaginous padding between these discs. These observations provided additional evidence for a lack of flexibility in plesiosaur necks.[36]
Everhart argued contrary to Carpenter's 1996 paper that polycotylids were present throughout the life of the Western Interior Seaway.[48]
Everhart finally described the partially digested partial plesiosaur skull he discovered in 1992. These were among the earliest known plesiosaur fossils in the Smoky Hill Chalk. He has since concluded that the animal that partially digested the remains was probably a shark, which would go on to vomit them up before they were buried and preserved.[77]
Michael Everhart found Charles H. Sternberg's account of the discovery of the Elasmosaurus sternbergi type specimen in his 1932 book. This allowed Everhart to verify the specimen's geographic and stratigraphic provenance.[104]
Everhart argued that the greater abundance of arc shaped marks and rounded divots in plesiosaur gastroliths compared to rocks deposited by ancient rivers and sea shores was evidence for their use in the breakdown of plesiosaurs' food.[97]
Everhart redescribed the Tylosaurus specimen USNM 8898 and its polycotylid dinner USNM 9468. Contrary to Sternberg's original assessment of the stomach contents as representing a "huge plesiosaur" Everhart found it to be a young polycotylid only about 2-2.5 m long.[55]
F. Robin O'Keefe and Luis M. Chiappe concluded the debate on plesiosaur reproduction, reporting the discovery of a gravid female plesiosaur with a single large embryo preserved inside her. They concluded that like marine mammals, but unlike many reptiles, plesiosaurs had a K-selected reproductive strategy.[10]
A reassessment of fossils attributed to the genus Polyptychodon is published by Madzia (2016), who considers the type species of this genus, P. interruptus, to be nomen dubium, and the genus Polyptychodon to be a wastebasket taxon.[134]
A redescription of the holotype specimen of Brancasaurus brancai and a study on the phylogenetic relationships of the species is published by Sachs, Hornung & Kear (2016), who consider the species Gronausaurus wegneri to be a junior synonym of B. brancai.[136]
A study on the mechanisms generating vertebral counts and their regionalisation during embryo development that were responsible for high plasticity of the body plan of sauropterygians is published by Soul & Benson (2017).[137]
A study on the function of the long neck in plesiosaurs as indicated by the anatomy of the neck is published by Noè, Taylor & Gómez-Pérez (2017).[138]
A study on the large, paired openings in the neck vertebrae of plesiosaurs and their implications for inferring the anatomy of the vascular system in the neck of plesiosaurs is published by Wintrich, Scaal & Sander (2017).[139]
A study on the swimming method of plesiosaurs is published by Muscutt et al. (2017).[140]
An assessment of the completeness of the plesiosaur fossil record is published by Tutin & Butler (2017).[141]
A study on the tooth formation cycle in elasmosaurid plesiosaurs is published by Kear et al. (2017).[143]
A redescription of the holotype specimen of Tuarangisaurus keyesi and a study on the phylogenetic relationships of the species is published by O'Gorman et al. (2017).[144]
A study on the anatomy of the vertebra of Vegasaurus molyi and its implications for the anatomy of the nervous system of the species is published by O'Gorman & Fernandez (2017).[145]
A study aiming to estimate metabolic rates and bone growth rates in eosauropterygians, especially in plesiosaurs, is published by Fleischle, Wintrich & Sander (2018).[153]
A study on the variability of the skull morphology in Simosaurus gaillardoti is published by de Miguel Chaves, Ortega & Pérez-García (2018).[154]
Morphologically diverse pliosaurid teeth are described from the Upper Jurassic (Tithonian) of the Kheta river basin (Eastern Siberia, Russia) and from the Lower Cretaceous (Berriasian and Valanginian) of the Volga region (European Russia) by Zverkov et al. (2018), who argue that their findings challenge the hypothesis that only one lineage of pliosaurids crossed the Jurassic–Cretaceous boundary.[157]
A study on a specimen of Cryptoclidus eurymerus from the Middle Jurassic (Callovian) of Peterborough (United Kingdom), with the left forelimb injured by a predator causing the loss of use of this limb but which nevertheless survived for some time after that injury, is published by Rothschild, Clark & Clark (2018), who also evaluate the implications of this specimen for the various hypotheses on plesiosaur propulsion.[160]
A study on the range of motion of the neck of an exceptionally preserved specimen of Nichollssaura borealis is published by Nagesan, Henderson & Anderson (2018).[161]
A study on the morphology of Thililua longicollis and on the phylogenetic relationships of members of the family Polycotylidae is published by Fischer et al. (2018), who name a new clade Occultonectia.[162]
An exceptionally well-preserved elasmosaurid basicranium, providing new information on the anatomy of the skull of elasmosaurids, is described from the Upper Cretaceous (lower Campanian) Rybushka Formation (Russia) by Zverkov, Averianov & Popov (2018).[165]
Redescription of Aristonectes quiriquinensis, providing new information on the anatomy of this species, is published by Otero, Soto-Acuña & O'keefe (2018).[166]
New elasmosaurid specimen is described from the upper Maatrichtian horizons of the Lopez de Bertodano Formation (Antarctica) by O’Gorman et al. (2018), representing one of the youngest non-aristonectine weddellonectian elasmosaurid specimens from Antarctica reported so far, documenting the presence of at least two different non-aristonectine elasmosaurids in Antarctica during the late Maastrichtian, and confirming the coexistence of aristonectine and non-aristonectine elasmosaurids in Antarctica until the end of the Cretaceous.[168]
Redescription of the holotype of Styxosaurus snowii and a study on the phylogenetic relationships of this species is published by Sachs, Lindgren & Kear (2018).[169]
Pathological fusions of neck vertebrae are reported in four plesiosaur specimens from different geological horizons by Sassoon (2019).[170]
A study on the morphology of the teeth and skull of Megacephalosaurus eulerti, and on their implications for assessing the phylogenetic relationships of this species, will be published by Madzia, Sachs & Lindgren (2019).[171]
A study on the skull morphology of two specimens of Dolichorhynchops bonneri from the Pierre Shale of South Dakota, as well as on the phylogenetic relationships of this species, is published by Morgan & O'Keefe (2019).[173]
A study on bone histology and ontogeny of the gravid specimen of Polycotylus latipinnus displayed at the Los Angeles County Museum of Natural History, and on its implications for interpreting a histological growth series in Dolichorhynchops bonneri, is published by O’Keefe et al. (2019).[174]
Skull and neck bones of an elasmosaurid plesiosaur are described from the CenomanianHegushi Formation (Japan) by Utsunomiya (2019), representing the oldest confirmed elasmosaurid in Japan and in East Asia.[175]
Sachs, Eggmaier and Madzia described a new species and genus of plesiosauroid plesiosaur, Franconiasaurus brevispinus, from the Toarcian deposits of Germany.[182]
^For the mythical creatures as Thunder Birds and Water Monsters, see Mayor (2005); "The Stone Medicine Bone, Pawnee Territory", page 178. For plesiosaurs as a specific source of these legends, see "Cheyenne Fossil Knowledge", page 211.
^José P. O'Gorman; Rodrigo A. Otero; Norton Hiller; John Simes; Marianna Terezow (2017). "Redescription of Tuarangisaurus keyesi (Sauropterygia; Elasmosauridae), a key species from the uppermost Cretaceous of the Weddellian Province: Internal skull anatomy and phylogenetic position". Cretaceous Research. 71: 118–136. Bibcode:2017CrRes..71..118O. doi:10.1016/j.cretres.2016.11.014.
^F. Robin O'Keefe; Rodrigo A. Otero; Sergio Soto-Acuña; Jose P. O'gorman; Stephen J. Godfrey; Sankar Chatterjee (2017). "Cranial anatomy of Morturneria seymourensis from Antarctica, and the evolution of filter feeding in plesiosaurs of the Austral Late Cretaceous". Journal of Vertebrate Paleontology. 37 (4): e1347570. Bibcode:2017JVPal..37E7570O. doi:10.1080/02724634.2017.1347570. hdl:11336/75235. S2CID91144814.
^Norton Hiller; José P. O’Gorman; Rodrigo A. Otero; Al A. Mannering (2017). "A reappraisal of the Late Cretaceous Weddellian plesiosaur genus Mauisaurus Hector, 1874". New Zealand Journal of Geology and Geophysics. 60 (2): 112–128. Bibcode:2017NZJGG..60..112H. doi:10.1080/00288306.2017.1281317. S2CID132037930.
^Sven Sachs; Benjamin P. Kear (2017). "Redescription of the elasmosaurid plesiosaurian Libonectes atlasense from the Upper Cretaceous of Morocco". Cretaceous Research. 74: 205–222. Bibcode:2017CrRes..74..205S. doi:10.1016/j.cretres.2017.02.017.
^José P. O’Gorman; Rodolfo A. Coria (2017). "A new elasmosaurid specimen from the upper Maastrichtian of Antarctica: new evidence of a monophyletic group of Weddellian elasmosaurids". Alcheringa: An Australasian Journal of Palaeontology. 41 (2): 240–249. Bibcode:2017Alch...41..240O. doi:10.1080/03115518.2016.1224318. hdl:11336/79006. S2CID132473041.
^Jose P. O’Gorman; Soledad Gouiric-Cavalli; Roberto A. Scasso; Marcelo Reguero; Juan J. Moly; Leonel Acosta-Burlaille (2018). "A Late Jurassic plesiosaur in Antarctica: Evidence of the dispersion of marine fauna through the Trans-Erythraean Seaway?". Comptes Rendus Palevol. 17 (3): 158–165. Bibcode:2018CRPal..17..158O. doi:10.1016/j.crpv.2017.10.005. hdl:11336/94465.
^Timothy Holland (2018). "The mandible of Kronosaurus queenslandicus Longman, 1924 (Pliosauridae, Brachaucheniinae), from the Lower Cretaceous of Northwest Queensland, Australia". Journal of Vertebrate Paleontology. 38 (5): e1511569. Bibcode:2018JVPal..38E1569H. doi:10.1080/02724634.2018.1511569. S2CID91599158.
^Rémi Allemand; Nathalie Bardet; Alexandra Houssaye; Peggy Vincent (2018). "New plesiosaurian specimens (Reptilia, Plesiosauria) from the Upper Cretaceous (Turonian) of Goulmima (Southern Morocco)". Cretaceous Research. 82: 83–98. Bibcode:2018CrRes..82...83A. doi:10.1016/j.cretres.2017.09.017.
^Rodrigo A. Otero; José P. O'Gorman; William L. Moisley; Marianna Terezow; Joseph McKee (2018). "A juvenile Tuarangisaurus keyesi Wiffen and Moisley 1986 (Plesiosauria, Elasmosauridae) from the Upper Cretaceous of New Zealand, with remarks on its skull ontogeny". Cretaceous Research. 85: 214–231. Bibcode:2018CrRes..85..214O. doi:10.1016/j.cretres.2017.09.007. hdl:11336/99631.
^Nikolay G. Zverkov; Alexander O. Averianov; Evgeny V. Popov (2018). "Basicranium of an elasmosaurid plesiosaur from the Campanian of European Russia". Alcheringa: An Australasian Journal of Palaeontology. 42 (4): 528–542. Bibcode:2018Alch...42..528Z. doi:10.1080/03115518.2017.1302508. S2CID132125319.
^Rodrigo A. Otero; Sergio Soto-Acuña; Frank R. O'keefe (2018). "Osteology of Aristonectes quiriquinensis (Elasmosauridae, Aristonectinae) from the upper Maastrichtian of central Chile". Journal of Vertebrate Paleontology. 38 (1): e1408638. Bibcode:2018JVPal..38E8638O. doi:10.1080/02724634.2017.1408638. S2CID90977078.
^Daniel Madzia; Sven Sachs; Johan Lindgren (2019). "Morphological and phylogenetic aspects of the dentition of Megacephalosaurus eulerti, a pliosaurid from the Turonian of Kansas, USA, with remarks on the cranial anatomy of the taxon". Geological Magazine. 156 (7): 1201–1216. Bibcode:2019GeoM..156.1201M. doi:10.1017/S0016756818000523. S2CID133859507.
^J.M. Quesada; A. Pérez-García; J.M. Gasulla; F. Ortega (2019). "Plesiosauria remains from the Barremian of Morella (Castellón, Spain) and first identification of Leptocleididae in the Iberian record". Cretaceous Research. 94: 8–24. Bibcode:2019CrRes..94....8Q. doi:10.1016/j.cretres.2018.10.010. S2CID134139253.
^Donald J. Morgan III; F. Robin O'Keefe (2019). "The cranial osteology of two specimens of Dolichorhynchops bonneri (Plesiosauria, Polycotylidae) from the Campanian of South Dakota, and a cladistic analysis of the Polycotylidae". Cretaceous Research. 96: 149–171. Bibcode:2019CrRes..96..149M. doi:10.1016/j.cretres.2018.11.027. S2CID134887820.
^Satoshi Utsunomiya (2019). "Oldest Elasmosauridae(Plesiosauria) in East Asia from the Upper Cretaceous Goshoura Group, Shishijima Island, southwestern Japan". Bulletin of the Osaka Museum of Natural History. 73: 23–35. doi:10.20643/00001333.
Roger B. J. Benson; Hilary F. Ketchum; Darren Naish & Langan E. Turner (2013). "A new leptocleidid (Sauropterygia, Plesiosauria) from the Vectis Formation (Early Barremian–early Aptian; Early Cretaceous) of the Isle of Wight and the evolution of Leptocleididae, a controversial clade". Journal of Systematic Palaeontology. 11 (2): 233–250. Bibcode:2013JSPal..11..233B. doi:10.1080/14772019.2011.634444. S2CID18562271.
Brown, B. (1913). "A new plesiosaur, Leurospondylus, from the Edmonton Cretaceous of Alberta". Bulletin of the American Museum of Natural History. 32: 606–615.
Buckland, W. (1837). Geology and Mineralogy, Considered With Reference to Natural Theology. London: William Pickering. p. 605.
Andrea Cau & Federico Fanti (2015). "High evolutionary rates and the origin of the Rosso Ammonitico Veronese Formation (Middle-Upper Jurassic of Italy) reptiles". Historical Biology: An International Journal of Paleobiology. 28 (7): 1–11. doi:10.1080/08912963.2015.1073726. S2CID86528030.
Conybeare, W. D. (1824). "On the discovery of an almost perfect skeleton of the Plesiosaurus". Transactions of the Geological Society of London. 2: 382–389.
Cruickshank, A. R. I.; Small, P. G.; Taylor, M. A. (1991). "Dorsal nostrils and hydrodynamically driven underwater olfaction in plesiosaurs". Nature. 352 (6330): 62–64. Bibcode:1991Natur.352...62C. doi:10.1038/352062a0. S2CID4353612.
Dames, W. (1895). "Die Plesiosaurier der s'u'ddeutschen Liasformation". Abhandlungen der Königlichen Preussischen Akademie der Wissenschaften: 1–83.
De la Beche, H. T.; Conybeare, W. D. (1821). "Notice of the discovery of a new fossil animal, forming a link between the Ichthyosaurus and the crocodile, together with general remarks on the osteology of Ichthyosaurus". Transactions of the Geological Society of London. 5: 559–594.
Ellis, Richard (2003). Sea Dragons - Predators of the Prehistoric Oceans. University Press of Kansas. ISBN978-0-7006-1269-7.
Everhart, Michael J. (2005). Oceans Of Kansas: A Natural History Of The Western Interior Sea. Life of the Past. Bloomington: Indiana University Press. p. 322.
Oliver Hampe (2013). "The forgotten remains of a leptocleidid plesiosaur (Sauropterygia: Plesiosauroidea) from the Early Cretaceous of Gronau (Münsterland, Westphalia, Germany)". Paläontologische Zeitschrift. 87 (4): 473–491. Bibcode:2013PalZ...87..473H. doi:10.1007/s12542-013-0175-3. S2CID129834688.
Huene, F. R. F. von (1923). "Ein neuer Plesiosaurier aus dem oberen Lias W'u'rttembergs". Jareshefte das Vereins F'u'r Vaterlandische Naturkunde in W'u'rttemberg. 79: 1–21.
Tai Kubo; Mark T. Mitchell & Donald M. Henderson (2012). "Albertonectes vanderveldei, a new elasmosaur (Reptilia, Sauropterygia) from the Upper Cretaceous of Alberta". Journal of Vertebrate Paleontology. 32 (3): 557–572. Bibcode:2012JVPal..32..557K. doi:10.1080/02724634.2012.658124. S2CID129500470.
Lydekker, R. (1889). Catalogue of the fossil Reptilia and Amphibia in the British Museum (Natural History). Part II. Containing the orders Ichthyopterygia and Sauropterygia. London: British Museum (Natural History). p. 307.
Lydekker, R. (1890). Catalogue of the fossil Reptilia and Amphibia in the British Museum (Natural History). Part IV. The orders Anomodontia, Ecaudata, Caudata, and Labyrinthodontia; and supplement. London: British Museum (Natural History). p. 295.
M-R
Mansel-Pleydell, J. C. (1888). "Fossil reptiles of Dorset". Proceedings of the Dorset Natural History and Antiquarian Field Club. 9: 1–40.
Mayor, Adrienne (2005). Fossil Legends of the First Americans. Princeton University Press. ISBN978-0-691-11345-6.
Rebecca Schmeisser McKean (2012). "A new species of polycotylid plesiosaur (Reptilia: Sauropterygia) from the Lower Turonian of Utah: extending the stratigraphic range of Dolichorhynchops". Cretaceous Research. 34: 184–199. Bibcode:2012CrRes..34..184S. doi:10.1016/j.cretres.2011.10.017.
José P. O’Gorman; Leonardo Salgado; Eduardo B. Olivero & Sergio A. Marenssi (2015). "Vegasaurus molyi, gen. et sp. nov. (Plesiosauria, Elasmosauridae), from the Cape Lamb Member (lower Maastrichtian) of the Snow Hill Island Formation, Vega Island, Antarctica, and remarks on Wedellian Elasmosauridae". Journal of Vertebrate Paleontology. 35 (3): e931285. Bibcode:2015JVPal..35E1285O. doi:10.1080/02724634.2014.931285. hdl:11336/53416. S2CID128965534.
Osborn, H. F. (1903). "The reptilian subclasses Diapsida and Synapsida and the early history of the Diaptosauria". Memoir of the American Museum of Natural History. 1: 449–507.
Owen, R. (1840a). "Report on British fossil reptiles. Part I.". Report of the British Association for the Advancement of Science 1839: 43–126.
Owen, R. (1840b). "A description of a specimen of the Plesiosaurus macrocephalus, Conybeare, in the collection of Viscount Cole, MP, DCL, FGS, &c". Transactions of the Geological Society of London. 5: 559–594.
Owen, R. (1860). "On the orders of fossil and Recent Reptilia, and their distribution in time". Report of the British Association for the Advancement of Science. 29: 153–166.
Owen, R. (1865). A monograph on the fossil Reptilia of the Liassic formations. Part I, Sauropterygia. Palaeontographical Society Monograph. Vol. 17. pp. 1–40.
Parkinson, J. (1822). Outline of Oryctology: An Introduction to the Study of Fossil Organic Remains; Especially of Those Found in British Strata. London. p. 350.
S-Z
Sauvage, H.-E. (1882). "Recherches sur les reptiles trouvé dans le Gault de l'est du Bassin de Paris". Mémoires de la Société Géologique de France. 2: 24–28.
Sauvage, H.-E. (1898). "Les reptiles et les poissons des terrains Mésozoîques du Portugal". Bulletin de la Société Géologique de France. 26: 442–446.
Schumacher, B. A.; Carpenter, K.; Everhart, M. J. (2013). "A new Cretaceous Pliosaurid (Reptilia, Plesiosauria) from the Carlile Shale (middle Turonian) of Russell County, Kansas". Journal of Vertebrate Paleontology. 33 (3): 613. Bibcode:2013JVPal..33..613S. doi:10.1080/02724634.2013.722576. S2CID130165209.
Sciau, J.; Crochet, J.-Y.; Mattei, J. (1990). "Le premier squelette de plesiosaure de France sur le Causse du Larzac (Toarcien, Jurassique Infe'rieur)". Geobios. 23 (1): 111–116. Bibcode:1990Geobi..23..111S. doi:10.1016/0016-6995(90)80021-7.
A. G. Sennikov; M. S. Arkhangelsky (2010). "On a Typical Jurassic Sauropterygian from the Upper Triassic of Wilczek Land (Franz Josef Land, Arctic Russia)". Paleontological Journal. 44 (5): 567–572. Bibcode:2010PalJ...44..567S. doi:10.1134/S0031030110050126. S2CID88505507.
Leonie Schwermann & Martin Sander (2011). Osteologie und Phylogenie von Westphaliasaurus simonsensii: Ein neuer Plesiosauride (Sauropterygia) aus dem Unteren Jura (Pliensbachium) von Sommersell (Kreis Höxter), Nordrhein-Westfalen, Deutschland [=Osteology and Phylogeny of Westphaliasaurus simonsensii, a new plesiosaurid (Sauropterygia) from the Lower Jurassic (Pliensbachian) of Sommersell (Höxter district), North Rhine-Westphalia, Germany]. Vol. 79. pp. 56 pp. ISBN978-3-940726-14-8. {{cite book}}: |journal= ignored (help)
Storrs, G. W. (1999). "An examination of Plesiosauria (Diapsida: Sauropterygia) from the Niobrara Chalk (upper Cretaceous) of central North America". Univ. Kansas Paleont. Cont. New Series. 11: 15.
Peggy Vincent & Roger B. J. Benson (2012). "Anningasaura, a basal plesiosaurian (Reptilia, Plesiosauria) from the Lower Jurassic of Lyme Regis, United Kingdom". Journal of Vertebrate Paleontology. 32 (5): 1049–1063. Bibcode:2012JVPal..32.1049V. doi:10.1080/02724634.2012.686467. S2CID86547069.
Peggy Vincent; Nathalie Bardet; Xabier Pereda Suberbiola; Baâdi Bouya; Mbarek Amaghzaz & Saïd Meslouh (2011). "Zarafasaura oceanis, a new elasmosaurid (Reptilia: Sauropterygia) from the Maastrichtian Phosphates of Morocco and the palaeobiogeography of latest Cretaceous plesiosaurs". Gondwana Research. 19 (4): 1062–1073. Bibcode:2011GondR..19.1062V. doi:10.1016/j.gr.2010.10.005.[dead link]
Watson, D. M. S. (1909). "A preliminary note on two new genera of Upper Liassic plesiosaurs". Memoirs of the Manchester Museum. 54: 1–26.
Welles, S. P. (1943). "Elasmosaurid plesiosaurs with description of new material from California and Colorado". Memoir of the University of California. 13: 125–215.
Welles, S. P. (1952). "A review of North American Cretaceous elasmosaurs". University of California Publications in the Geological Sciences. 29: 47–144.
Welles, S. P. (1962). "A new species of elasmosaur from the Aptian of Columbia and a review of the Cretaceous plesiosaurs". University of California Publications in the Geological Sciences. 44: 1–96.
Winkler, T. C. (1873). "Le Plesiosaurus dolichodeirus, Conyb., du Musée Teyler". Archives du Musée Teyler. 3: 219–233.
Woodward, H. B. (1893). The Jurassic rocks of Britain. Volume III. The Lias of England and Wales (Yorkshire excepted). Memoir of the Geological Survey of the United Kingdom. London. p. 399.{{cite book}}: CS1 maint: location missing publisher (link)
External links
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