Elasmosaurus

***Elasmosaurus*** (meaning "thin-plate lizard) 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) long, *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.Measuring 10.3 meters (34 ft) long, *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.

## Description

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; 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. 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. In 1952 the American paleontologist Samuel Welles estimated the body length to have been 10.3 meters (34 ft). 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). The living animal would have been slightly larger due to cartilage present between the vertebral bodies, and was estimated by Cope at roughly 13.7 meters (45 ft).

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.

The pectoral and pelvic girdles of the holotype specimen were noted as missing by 1906, but observations about these elements were since made based on the original descriptions and figures from the late 19th century. The shoulder blades (scapulae) were fused and met at the midline, bearing no trace of a median bar. The upper processes of the shoulder blades were very broad, and the "necks" of the shoulder blades were long. The pectoral girdle had a long bar at the middle, a supposedly advanced feature thought to be absent from juvenile plesiosaurs. The ischia (a pair of bones that formed part of the pelvis) were joined at the middle, so that a medial bar was present along the length of the pelvis, a feature usually not found in plesiosaurs. Like other elasmosaurids (and plesiosaurs in general), *Elasmosaurus* would have had large, paddle-like limbs with very long digits. The paddles at the front (the pectoral paddles) were longer than those at the back (the pelvic paddles).

### Vertebrae

Unlike those of many other long-necked animals, the individual neck vertebrae were not particularly elongated; rather, the extreme neck length was achieved by a much increased number of vertebrae. *Elasmosaurus* differed from all other plesiosaurs by having 72 neck (or cervical) vertebrae; more may have been present but were later lost to erosion or after excavation. Only *Albertonectes* had more neck vertebrae, 75, and the two are the only plesiosaurs with a count higher than 70; more than 60 vertebrae is very derived (or "advanced") for plesiosaurs.

The atlas and axis bone complex, consisting of the first two neck vertebrae and articulated with the back of the skull, was long, low, and horizontally rectangular in side-view. The centra, or "bodies", of these vertebrae were co-ossified in the holotype specimen, which indicates it was an adult. The neural arches of these vertebrae were very thin and rather high, which gave the neural canal (the opening through the middle of the vertebrae) a triangular outline when seen from the back. The lower part of the neural canal was narrow towards the back by the axis, where it was half the breadth of the centrum. It became broader towards the front, where it was almost the same breadth as the centrum of the atlas. The neural arches were also more robust there than in the axis, and the neural canal was higher. The neural spine was low and directed upwards and back. The centra of the atlas and axis were of equal length, and had a quadratic shape in side view. The surface (or facet) where the axis articulated with the next vertebra had an oval outline, and an excavation for the neural canal in the middle of its upper edge. A distinct keel ran along the lower middle of the atlas and axis vertebrae.

Most of the neck vertebrae were compressed sideways, especially at the middle of the neck. A crest (also termed ridge or keel) ran longitudinally along the side of the neck vertebrae (a feature typical of elasmosaurids), visible from the third to the fifty-fifth vertebrae, at the hind part of the neck. This crest was positioned at the middle of the centrum in the front vertebrae, and at the upper half of the centrum from the 19th vertebra and onwards. The crest would have served to anchor the musculature of the neck. The centra differed in shape depending on the position of the vertebrae in the neck; that of the third vertebra was about as long as it was broad, but the centra became longer than broad from the fourth vertebra and onwards. The centra became more elongated at the middle of the neck, but became shorter again at the back of the neck, with the length and breadth being about equal at the 61st vertebra, and those of the hindmost vertebrae being broader than long. The articular surfaces of the vertebrae in the front of the neck were broad oval, and moderately deepened, with rounded, thickened edges, with an excavation (or cavity) at the upper and lower sides. Further back in the front part of the neck, around the 25th vertebra, the lower edge of the articular facets became more concave, and the facet shaped like a quadrate with rounded edges. By the 63rd vertebra, the articular facet was also quadratic in shape with rounded edges, whereas the centra of the hindmost vertebrae had a broad oval outline.

The neural arches of the neck vertebrae were well fused to the centra, leaving no visible sutures, and the neural canal was narrow in the front vertebrae, becoming more prominently developed in the hind vertebrae, where it was as broad as high, and almost circular. The pre-and post-zygapophyses of the neck vertebrae, processes that articulated adjacent vertebrae so they fit together, were of equal length; the former reached entirely over the level of the centrum whereas the latter reached only with their back half. The neural spines of the neck vertebrae appear to have been low, and almost semi-circular by the 20th vertebra. The facets where the neck ribs articulated with the neck vertebrae were placed on the lower sides of the centra, but were only placed higher in the last three vertebrae, reaching around the middle of the sides. The neck ribs were semicircular to quadratic in side view, and were directed rather straight down. The bottom of each neck vertebrae had pairs of nutritive foramina (openings) at the middle, separated by a ridge, which became progressively more prominent and thickened towards the back of the neck.

The vertebrae that transitioned between the neck and back (or dorsal) vertebrae in the pectoral region of plesiosaurs, close to the front margin of the forelimb girdle, are often termed pectoral vertebrae. *Elasmosaurus* had three pectoral vertebrae, which is a common number for elasmosaurids. The rib facets of the pectoral vertebrae were triangular in shape and situated on transverse processes, and the centra bore pairs of nutritive foramina in the middle of the lower sides. The back vertebrae had rib facets level with the neural canal, and the front and back part of the transverse processes here had distinct ridges on their margins. Here the rib facets where placed higher than the transverse processes, separating the two, and were oval to rectangular in outline. The pre-zygapophyses here were shorter than those in the neck and pectoral vertebrae, and only reached above the level of the centrum with the front third of their length. The post-zygapophyses reached over the level of the centrum with the back half of their length. Back vertebrae are not useful for distinguishing between elasmosaurids, since they are not diagnostic at the genus level.

*Elasmosaurus* had four sacral vertebrae (the fused vertebrae that form the sacrum connected to the pelvis), a number typical of elasmosaurids. The transverse processes here were very short, and the rib facets increased in size from the first to the fourth sacral vertebra. A ridge ran along the top of these vertebrae, and the lower sides of the centra were rounded, and bore pairs of nutritive foramina, separated by low ridges. The first tail (or caudal) vertebra could be distinguished by the preceding sacral vertebra by having smaller rib facets, and by being positioned in the lower half of the centrum. These vertebrae were almost circular in shape, and the first two bore a narrow keel in the middle of the upper side. The rib facets of the tail vertebrae were located on the lower side of the centra, and their oval shape became larger and broader from the third vertebra and onwards, but became smaller from the 14th vertebra. Here, the pre-zygapophyses also reached over the level of the centra for most of their length, while the post-zygapophyses reached over this level by half their length. The lower part of the centra were rounded from the first to the third tail vertebrae, but concave from the fourth to the 18th. The usual number of tail vertebrae in elasmosaurids is 30. Since the last tail-vertebrae of elasmosaurids were fused into a structure similar to the pygostyle of birds, it is possible this supported a tail-fin, but the shape it would have had is unknown.

## *Elasmosaurus* in The Amazing Dinosaur Adventure

### Anonymous

*Elasmosaurus* from the opening narration to *Journey to Big Water*