Description
Elasmosaurus (/ ɪˌlæzməˈsɔːrəs, -moʊ-/;[1] ) is a genus of plesiosaur that lived in North America during the Campanian stage of the Late Cretaceous period, about 80.5 million years ago. The first specimen was discovered in 1867 near Fort Wallace , Kansas, US, and was sent to the American paleontologist Edward Drinker Cope , who named it E. platyurus in 1868. The generic name means "thin-plate reptile", and the specific name means "flat-tailed". Cope originally reconstructed the skeleton of Elasmosaurus with the skull at the end of the tail, an error which was made light of by the paleontologist Othniel Charles Marsh , and became part of their "Bone Wars " rivalry. Only one incomplete Elasmosaurus skeleton is definitely known, consisting of a fragmentary skull, the spine, and the pectoral and pelvic girdles , and a single species is recognized today; other species are now considered invalid or have been moved to other genera.
Measuring 10.3 meters (34 ft) in length, Elasmosaurus would have had a streamlined body with paddle-like limbs, a short tail, a small head, and an extremely long neck. The neck alone was around 7.1 meters (23 ft) long. Along with its relative Albertonectes , it was one of the longest-necked animals to have lived, with the second largest number of neck vertebrae known, 72, 4 less than Albertonectes. The skull would have been slender and triangular, with large, fang-like teeth at the front, and smaller teeth towards the back. It had six teeth in each premaxilla of the upper jaw, and may have had 14 teeth in the maxilla and 19 in the dentary of the lower jaw. Most of the neck vertebrae were compressed sideways, and bore a longitudinal crest or keel along the sides.
The family Elasmosauridae was based on the genus Elasmosaurus, the first recognized member of this group of long-necked plesiosaurs. Elasmosaurids were well adapted for aquatic life, and used their flippers for swimming. Contrary to earlier depictions, their necks were not very flexible, and could not be held high above the water surface. It is unknown what their long necks were used for, but they may have had a function in feeding. Elasmosaurids probably ate small fish and marine invertebrates , seizing them with their long teeth, and may have used gastroliths (stomach stones) to help digest their food. Elasmosaurus is known from the Pierre Shale formation, which represents marine deposits from the Western Interior Seaway .
In early 1867, the American army surgeon Theophilus Hunt Turner and the army scout William Comstock explored the rocks around Fort Wallace , Kansas , where they were stationed during the construction of the Union Pacific Railroad . Approximately 23 kilometers (14 mi) northeast of Fort Wallace, near McAllaster , Turner discovered the bones of a large fossil reptile in a ravine in the Pierre Shale formation, and though he had no paleontological experience, he recognized the remains as belonging to an "extinct monster". In June, Turner gave three fossil vertebrae to the American scientist John LeConte , a member of the railway survey, to take back east to be identified. In December, LeConte delivered some of the vertebrae to the American paleontologist Edward Drinker Cope at the Academy of Natural Sciences of Philadelphia (ANSP, known since 2011 as the Academy of Natural Sciences of Drexel University). Recognizing them as the remains of a plesiosaur , larger than any he had seen in Europe, Cope wrote to Turner asking him to deliver the rest of the specimen, at the ANSP's expense.[2] [3] [4]
In December 1867 Turner and others from Fort Wallace returned to the site and recovered much of the vertebral column, as well as concretions that contained other bones; the material had a combined weight of 360 kilograms (800 lb). The fossils were dug or pried out of the relatively soft shale with picks and shovels, loaded on a horse-drawn wagon, and transported back to Fort Wallace. Cope sent instructions on how to pack the bones, which were thereafter sent in hay-padded crates on a military wagon east to the railroad, which had not yet reached the fort. The specimen arrived in Philadelphia by rail in March 1868, whereafter Cope examined it hurriedly; he reported on it at the March ANSP meeting, during which he named it Elasmosaurus platyurus. The generic name Elasmosaurus means "thin-plate reptile", in reference to the "plate" bones of the sternal and pelvic regions, and the specific name platyurus means "flat-tailed", in reference to the compressed "tail" (actually the neck) and laminae of the vertebrae there.[3] [5] [6] [7] [8]
Cope requested that Turner search for more parts of the Elasmosaurus specimen, and was sent more fossils during August or September 1868. The ANSP thanked Turner for his "very valuable gift" at their meeting in December 1868, and Turner visited the museum during spring, at a time when Cope was absent. Turner died unexpectedly at Fort Wallace on July 27, 1869, without seeing the completion of the work he began, but Cope continued to write him, unaware of his death until 1870. The circumstances around Turner's discovery of the type specimen were not covered in Cope's report, and remained unknown until Turner's letters were published in 1987. Elasmosaurus was the first major fossil discovery in Kansas (and the largest from there at the time), and marked the beginning of a fossil collecting rush that sent thousands of fossils from Kansas to prominent museums on the American east coast.[3] Elasmosaurus was one of few plesiosaurs known from the New World at the time, and the first recognized member of the long-necked family of plesiosaurs, the Elasmosauridae .[2]
In 1869 Cope scientifically described and figured Elasmosaurus, and the preprint version of the manuscript contained a reconstruction of the skeleton which he had earlier presented during his report at an ANSP meeting in September 1868. The reconstruction showed Elasmosaurus with a short neck and a long tail, unlike other plesiosaurs, and Cope was also unsure whether it had hind limbs. At an ANSP meeting a year and a half later, in March 1870, the American paleontologist Joseph Leidy (Cope's mentor) noted that Cope's reconstruction of Elasmosaurus showed the skull at the wrong end of the vertebral column, at the end of the tail instead of the neck. Cope had apparently concluded that the tail vertebrae belonged to the neck, since the jaws had been found at that end of the skeleton, even though the opposite end terminated in the axis and atlas bones that are found in the neck. Leidy also concluded that Elasmosaurus was identical to Discosaurus , a plesiosaur he had named in 1851.[8] [9] [10] [3]
To hide his mistake, Cope attempted to recall all copies of the preprint article, and printed a corrected version with a new skeletal reconstruction that placed the head on the neck (though it reversed the orientation of the individual vertebrae) and different wording in 1870. In a reply to Leidy, Cope claimed that he had been misled by the fact that Leidy had arranged the vertebrae of Cimoliasaurus in the reverse order in his 1851 description of that genus, and pointed out that his reconstruction had been corrected. Cope also rejected the idea that Elasmosaurus and Discosaurus were identical, and noted that the latter and Cimoliasaurus did not have any distinguishing features. Though Cope had tried to destroy the preprints, one copy came to the attention of the American paleontologist Othniel Charles Marsh , who made light of the mistake. This led to antagonism between Cope, who was embarrassed by the mistake, and Marsh, who brought up the mistake repeatedly for decades. Marsh returned to the issue during their controversy in the New York Herald in the 1890s (Marsh claimed he had pointed out the error to Cope immediately), when their dispute gained widespread public attention. The argument was part of the "Bone Wars " rivalry between the two, and is well known in the history of paleontology.[3] [9] [11] [12] [13] [14]
Because of Cope's reputation as a brilliant paleontologist, it has been questioned why he would make such an obvious anatomical error. It has been suggested that, as a unique specimen in 1868, the original Elasmosaurus may have been hard to interpret based on the knowledge available at the time. Also, Cope initially thought it consisted of two specimens of different animals – in an 1868 letter to LeConte, Cope had referred to the supposed "smaller specimen" as Discosaurus carinatus. Cope was only in his late twenties and not formally trained in paleontology, and may have been influenced by Leidy's mistake of reversing the vertebral column of Cimoliasaurus. In 2002 the American art historian Jane P. Davidson noted that the fact that other scientists early on had pointed out Leidy's error argues against this explanation, adding that Cope was not convinced he had made a mistake. Plesiosaur anatomy was sufficiently well known at the time that Cope should not have made the mistake, according to Davidson.[9] Cope did little work on the specimen since his 1870 description, and it was kept in storage for nearly 30 years.[3] It was only redescribed in detail in 2005 by the German paleontologist Sven Sachs.[2]
Today, the incomplete holotype specimen , cataloged as ANSP 10081, is the only definite specimen of Elasmosaurus. It was long exhibited, but is now stored in a cabinet with other assigned fragments. The specimen consists of the premaxillae, part of the hind-section of the right maxilla, two maxilla fragments with teeth, the front part of the dentaries, three more jaw fragments, two cranial fragments of indeterminable identity, 72 neck vertebrae, including the atlas and axis, 3 pectoral vertebrae, 6 back vertebrae, 4 sacral vertebrae, 18 tail vertebrae, as well as rib fragments.[15] [2] In 2013 an incomplete neck vertebra centrum of the holotype that had been mentioned by Cope but thought to have been lost was rediscovered in storage by Sachs, and the neck vertebra count was revised from 71 to 72.[15] In 1986 a three-dimensional reconstruction of the holotype skeleton was completed and is now displayed at the ANSP. This cast was later copied by the company Triebold Paleontology Incorporated , and replicas were provided to other museums. The replica at the Fort Wallace Museum measures about 12.8 meters (42 ft) in length.[3]
Though Cope described and figured the pectoral and pelvic girdles of Elasmosaurus in 1869 and 1875, these elements were noted as missing from the collection by the American paleontologist Samuel Wendell Williston in 1906. Cope had loaned these elements to the English sculptor Benjamin Waterhouse Hawkins to help prepare them out of their surrounding concretions. At the time, Hawkins was working on a "Paleozoic Museum " in New York's Central Park , where a reconstruction of Elasmosaurus was to appear, an American equivalent to his life-sized Crystal Palace Dinosaurs in London. In May 1871 much of the exhibit material in Hawkins' workshop was destroyed by vandals for unclear reasons and their fragments buried; it is possible that the girdle elements of Elasmosaurus were at the workshop and were likewise destroyed. Nothing was subsequently mentioned about their loss by Hawkins or Cope.[2] [3] [16] [17] [18] In 2018, Davidson and Everhart documented the events leading up to the disappearance of these fossils, and suggested that a photo and drawing of Waterhouse's workshop from 1869 appear to show concretions on the floor that may have been the unprepared girdles of Elasmosaurus. They also noted that conceptual sketches of the Palaeozoic Museum show that the model Elasmosaurus was originally envisioned with a long "tail", though later updated with a long neck. Davidson and Everhart concluded that the girdle fossils were most likely destroyed in Hawkins' workshop.[17]
Fossils that may have belonged to the holotype were found by the American geologist Benjamin Franklin Mudge in 1871, but have probably been lost since.[3] Additional plesiosaur fossils were recovered near the original locality in 1954, 1991, 1994, and 1998, including back vertebrae, ribs, gastralia (belly ribs), and gastroliths . As none of these elements overlap with those of the holotype specimen, in 2005 the American paleontologist Michael J. Everhart concluded they belonged to the same individual, and that the parts had been separated before burial of the carcass. He also noted that a small stone wedged in the neural canal of one of the tail vertebrae of the holotype may be a gastrolith, based on its polished appearance.[19] In 2007 the Colombian paleontologists Leslie Noè and Marcela Gómez-Pérez expressed doubt that the additional elements belonged to the type specimen, or even to Elasmosaurus, due to lack of evidence. They explained that elements missing from the holotype may have been lost to weathering or simply not collected, and that parts may have been lost or damaged during transportation or preparation. Gastroliths may also not have been recognized as such during collection, since such stones were not reported from a plesiosaur until ten years after.[20]
In 2017 Sachs and Joachim Ladwig suggested that a fragmentary elasmosaurid skeleton from the upper Campanian of Kronsmoor in Schleswig-Holstein , Germany, and housed in the Naturkunde-Museum Bielefeld , may have belonged to Elasmosaurus. Additional parts of the same skeleton are housed at the Institute for Geology of the University of Hamburg , as well as in private collections. Combined, the specimen consists of neck, back and tail vertebrae, phalanges , a tooth, limb elements, 110 gastroliths, and unidentifiable fragments.[21]
Though the only known specimen of Elasmosaurus (holotype specimen ANSP 10081) is fragmentary and missing many elements, related elasmosaurids show it would have had a compact, streamlined body, long, paddle-like limbs, a short tail, a proportionately small head, and an extremely long neck. The neck of Elasmosaurus is estimated at 7.1 meters (23 ft) in length;[22] thus, Elasmosaurus and its relative Albertonectes were some of the longest-necked animals ever to have lived, with the largest number of neck vertebrae of any known vertebrate animals.[23] [15] In spite of their many neck vertebrae, the necks of elasmosaurids were less than half as long as those of the longest-necked sauropod dinosaurs .[22] Initially, in his 1869 description of Elasmosaurus, Cope estimated the length of the animal by summing up vertebral lengths and estimations of missing parts, resulting in a total length of 13.1 meters (43 ft); he believed that the living animal would have been slightly larger due to cartilage present between the vertebral bodies, and was estimated at roughly 13.7 meters (45 ft).[8] However, in 1952, the American paleontologist Samuel Welles estimated the body length to have been 10.3 meters (34 ft).[24]
Like other elasmosaurids, Elasmosaurus would have had a slender, triangular skull. The snout was rounded and almost formed a semi-circle when viewed from above, and the premaxillae (which form the front of the upper jaw) bore a low keel at the midline. It is uncertain how many teeth Elasmosaurus had, due to the fragmentary state of the fossils. It probably had six teeth in each premaxilla, and the teeth preserved there were formed like large fangs. The number of premaxillary teeth distinguished Elasmosaurus from primitive plesiosauroids and most other elasmosaurids, which usually had fewer. The two teeth at the front were smaller than the succeeding ones, and were located between the first two teeth in the dentaries of the lower jaws. The known teeth of the front part of the lower jaw were large fangs, and the teeth at the back of the jaws appear to have been smaller. The dentition of elasmosaurids was generally heterodont (irregular throughout the jaws), with the teeth becoming progressively smaller from front to back. The maxillae (largest tooth bearing bone of the upper jaw) of elasmosaurids usually contained 14 teeth, whereas the dentaries (the main part of the lower jaws) usually contained 17 to 19. The teeth interlocked, and their tooth crowns were slender and rounded in cross-section. The mandibular symphysis (where the two halves of the lower jaw connected) was well ossified , with no visible suture .[23] [2]