''Muraenosaurus'' (from the Latin "''

Muraena ''Muraena'' is a genus of twelve species of large eels in the family Muraenidae. This genus is common in the Mediterranean, and is abundantly represented in tropical and subtropical seas, especially in rocky parts or on coral reefs. In the majo ...

''" meaning "eel" and "''Sauros''" meaning lizard) is an extinct genus of cryptoclidid

plesiosaur The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...

reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...

from the

Oxford Clay The Oxford Clay (or Oxford Clay Formation) is a Jurassic marine sedimentary rock formation underlying much of southeast England, from as far west as Dorset and as far north as Yorkshire. The Oxford Clay Formation dates to the Jurassic, specific ...

of Southern

England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...

. The genus was given its name due to the eel-like appearance of the long neck and small head. ''Muraenosaurus'' grew up to in length and lived roughly between 160 Ma (million years ago) and 164 Ma in the

Callovian In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 165.3 ± 1.1 Ma (million years ago) and 161.5 ± 1.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...

of the middle Jurassic. Charles E. Leeds collected the first ''Muraenosaurus'' which was then described by

H. G. Seeley Harry Govier Seeley (18 February 1839 – 8 January 1909) was a British paleontologist. Early life Seeley was born in London on 18 February 1839, the second son of Richard Hovill Seeley, a goldsmith, and his second wife Mary Govier. When his fa ...

.Seeley, HG. 1874. On ''Murænosaurus Leedsii'', a Plesiosaurian from the Oxford Clay. Part I. ''Quarterly Journal of the Geological Society'' 30: 197-208. The specimen may have suffered some damage due to the casual style of Charles Leeds’ collection. The first muraenosaur was recovered with pieces missing from the skull and many of the caudal vertebrae absent. Because the animal was described from Charles Leeds’ collection it was given the name ''Muraenosaurus Leedsii''. ''M. leedsii'' is the most complete specimen belonging to the genus ''Muraenosaurus'' and also the only species that is undoubtedly a member of the genus. Two other species have been tentatively referred to as members of the genus ''Muraenosaurus'': ''M. reedii'' and ''Muraenosaurus beloclis'' Seeley 1892,Wilhelm BC. 2010. A New Partial Skeleton of a Cryptocleidoid Plesiosaur from the Upper Jurassic Sundance Formation of Wyoming. '' Journal of Vertebrate Paleontology '', ''30'', 6, 1736-1742.Muraenosaurus? Reedii, Sp. Nov. and Tricleidus? Laramiensis Knight, American Jurassic Plesiosaurs. ''The Journal of Geology'', ''20'', 4, 344-352. which in 1910 became the separate genus ''

Picrocleidus ''Picrocleidus'' is an extinct genus of plesiosaur from the Middle Jurassic Oxford Clay Formation ( Callovian stage) of the United Kingdom. Discovery and naming The holotype was discovered in the Oxford Clay Formation in Peterborough, Eng ...

''. ''Muraenosaurus reedii'' was discovered in

Wyoming Wyoming ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States, Western United States. It borders Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho t ...

and described by Maurice Mehl of the University of Chicago. The fossilized fragments found consisted of pieces of the pectoral and pelvic girdles, several vertebrae, ribs, and a relatively complete left pectoral paddle. Connections between ''M. leedsii'' and ''M. reedii'' were drawn when examining the ribs structure and paddle of ''M. reedii''. Both had a similarly shaped, elongated humerus and relatively short phalanges. The species ''M. reedii'' is in fact a

junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...

of '' Pantosaurus''. ''Picrocleidus beloclis'' is another plesiosaur originating in the middle Jurassic and found in the Oxford clay formation. ''

'' has previously been considered synonymous with ''Muraenosaurus'' although there are doubts in the relationship between the two groupings.

Description

''Muraenosaurus'' was a medium-sized plesiosaur, with the largest specimen measuring up to long. The plesiosaur had 66 pre-sacral vertebrae; 44 of which were cervical. Starting from the base of the skull and moving posteriorly the

cervical vertebrae In tetrapods, cervical vertebrae (: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In saurop ...

elongate and the neural spines broaden. This may be indicative of increased muscle attachment towards the base of the neck.Richards, CD. 2011. ''Pleisiosaur body shape and its impact on hydrodynamic properties''. Huntington, WV: Marshal University Libraries There is also a gradual flattening and reduction of the zygapohesis until about the 30th vertebra where all traces of the platform are gone. Posterior to the 44 cervical vertebrae, ''Muraenosaurus'' had 3 pectoral vertebrae and 19

thoracic vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebra (anatomy), vertebrae of intermediate size between the ce ...

. The pectoral vertebrae resemble the terminal cervical vertebrae but display a broadened face of the centrum and a slight posterior incline in the neural spines. In the dorsal vertebrae the rib has been raised onto the

neural arch Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...

. The neural spines have elongated anteroposteriorly and compressed vertically. The

pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans, it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists o ...

in ''Muraenosaurus'' is broader than in most plesiosaurs and helped to situate the animal as a member of ''Cryptoclididae''. The coracoids reach widths of nearly 14 inches. The forelimb is compressed mediolaterally when compared to the hind limb as well as limbs of other plesiosaurs.O’Keefe FR. 2001. Ecomorphology of plesiosaur flipper geometry. ''Journal of Evolutionary Biology'', ''14'', 6, 987-991 This compression is represented in the

aspect ratio The aspect ratio of a geometry, geometric shape is the ratio of its sizes in different dimensions. For example, the aspect ratio of a rectangle is the ratio of its longer side to its shorter side—the ratio of width to height, when the rectangl ...

of muraenosaur limbs. The aspect ratio of the hind limbs is much greater than that of the fore limbs, representing a longer and more slender form. The high aspect ratio in the forelimbs may have been used for increasing maneuverability at some cost to the animal's endurance.

Classification

When

Harry Seeley Harry Govier Seeley (18 February 1839 – 8 January 1909) was a British paleontologist. Early life Seeley was born in London on 18 February 1839, the second son of Richard Hovill Seeley, a goldsmith, and his second wife Mary Govier. When his fa ...

first described ''Muraenosaurus'' it was thought to belong to

Elasmosauridae Elasmosauridae, often called elasmosaurs or elasmosaurids, is an extinct family of plesiosaurs that lived from the Hauterivian stage of the Early Cretaceous to the Maastrichtian stage of the Late Cretaceous period (c. 130 to 66 mya). The taxo ...

instead of

Cryptoclididae Cryptoclididae is a family of medium-sized plesiosaurs that existed from the Middle Jurassic to the Early Cretaceous. They had long necks, broad and short skulls and densely packed teeth. They fed on small soft-bodied preys such as small fish and ...

. Muraenosaurs and elasmosaurs share a plesiosauromorph body type typified by having a small head at the end of a long neck.O’Keefe, FR. 2001. A cladistics analysis and taxonomic revision of the ''Plesiosauria (Reptilia: Sauropterygia)''. ''Helsinki: Finnish Zoological and Botanical Pub. Board'' Like elasmosaurids, ''Muraenosaurus'' has a long neck. Of the 66 presacral

vertebrae Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...

in ''Muraenosaurus'', 44 of them are cervical. It was initially believed that this innovation leads to a largely flexible neck and a relatively short and sturdy body. More recent research has shown that while mobile, the neck of plesiosauromorphs was not as flexible as previously thought.Zammit M. 2008. Elasmosaur (Reptilia: Sauropterygia) neck flexibility: Implications for feeding strategies. ''Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology '', ''150'', 2, 124-130. The head of ''Muraenosaurus'' is also very small compared to both its neck and body length, measuring only about long.Andrews, CW. 1913. ''A descriptive catalogue of the Marine Reptiles of the Oxford Clay, Part II''. British Museum (Natural History). pp.205pp. Both of these traits are common in elasmosaurids which led to the initial diagnosis of muraenosaurs in the family ''Elasmosauridae''. However, this is actually a case of

convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...

between cryptoclidids and elasmosaurs.O’Keefe FR. 2002. The evolution of plesiosaur and pliosaur morphotypes in the Plesiosauria (Reptilia: Sauropterygia). ''Paleobiology'', ''28'', 1, 101-112 The clade of cryptoclidids creates a unique tree in relation to the paleomorphology of discovered specimens. For example, within cryptocleidoidae there are short necked plesiosaurs such as ''

Kimmerosaurus ''Kimmerosaurus'' ("lizard from Kimmeridge") is an extinct genus of plesiosaur from the family Cryptoclididae. It is known from remains found in England and Norway. Discovery There are very few fossil remains of ''Kimmerosaurus'' known. In fa ...

'' as well as the long necked ''Muraenosaurus''. The defining features of Cryptocleidoidia include a low fin aspect ratio, a wide rounded interpterygoid vacuity, and extreme specialization of the cheek region. The interpterygoid vacuity is completely absent in elasmosaur species but well represented in cryptoclidids, including ''Muraenosaurus''. The aspect ratio of both the fore and hind limbs of ''Muraenosaurus'' is far lower than the aspect ratio of elasmosaur specimens. The forelimbs especially resemble the flippers of

Pliosauridae Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous ( Rhaetian to Turonian stages). The family is more inclusive than the archetypal short-necked large headed species that are placed ...

, an extinct group of less elongate marine reptiles that shared the seas with plesiosaurs, more than they resemble the typical plesiosaur morphology. '' Cryptoclidus'', another cryptoclidid plesiosaurimorph, shares this trait with ''Muraenosaurus''.

Paleobiology

Habitat

''Muraenosaurus'' was initially discovered in the Oxford Clay which represents an ancient sea that was both shallow, with an average depth less than 50 meters, and warm (20 °C).Martill DM. 1994. The trophic structure of the biota of the Peterborough Member, Oxford Clay Formation (Jurassic), UK. ''Journal of the Geological Society '', ''151'', 1, 173-194.Martill DM. 1991. ''Fossils of the Oxford Clay''. Palaeontological Association, 1991. The Oxford Clay deposits are found in southern England and parts of France. The sea was abundant with nutrients and light filtered easily through the shallow water to create a highly productive ecosystem. The sea floor was littered with

bivalves Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...

arthropods Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...

gastropods Gastropods (), commonly known as slugs and snails, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and from the land. Ther ...

and

foraminifera Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...

while the pelagic zone was home to a wide variety of species from marine reptiles to teleosts. The Oxford Clay was so productive that over 100 genera have been recovered from the sediment.

Diet and predation

''Muraenosaurus''’ role in the ecosystem is probably more comparable to elasmosaurs than other cryptoclidids due to the plesiosauromorph body plans shared between elasmosaurs and muraenosaurs. Cryptoclidids have varying morphology and it is difficult to assess their ecological role as a collective unit. Long necked plesiosaurs have been discovered with varying contents lithified within their stomachs which give some indication of what ''Muraenosaurus'' may have been eating.McHenry CR. 2005. Bottom-feeding plesiosaurs. ''Science'', ''310'', 5745. Preferred prey types seem to be

teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...

fish and

cephalopods A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...

. In order to exploit such different prey species, plesiosaurs would need multiple different feeding strategies. Muraenosaurus leedsii

Initially, long necked plesiosaurs were thought to be strictly fish eaters due to their conical teeth, a shared trait with modern

piscivores A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...

. It was proposed that plesiosaurimophs would swim with a straight neck and attack

pelagic The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...

fishes. The straight neck would have been used in order to avoid creating drag by arching its neck upward into the water column. Additionally, some researchers have proposed that by swimming with its head directly in front of its body, plesiosaurs would be able to reach pelagic fishes before they felt the change in water pressure caused by the large body of the plesiosaur. Essentially, the head would precede the pressure difference. It is also proposed that species like ''Muraenosaurus'' fed upon benthic fishes by floating above them and reaching its head down into the benthos. Plesiosauromorphs may also have employed a strategy called benthic grazing where they would harvest relatively immobile species such as cephalopods from the sea floor. Gastroliths have often been found within the stomachs of extinct marine reptiles and have been associated with the plesiosauromorph body type. One of the proposed uses for gastroliths is the grinding of tough, shelled foods like cephalopods.Cicimurri DJ. 2001. An Elasmosaur with Stomach Contents and Gastroliths from the Pierre Shale (Late Cretaceous) of Kansas. ''Transactions of the Kansas Academy of Science '', ''104'', 3, 129-142. The food chain did not stop at the plesiosaurimorphs. The oceans in the

Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...

were dominated by pliosaurs,

apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...

s that would have preyed upon species like ''Muraenosaurus''. Pliosaurs were not the only animals that would have fed on ''Muraenosaurus'' though. Evidence of this comes from several cases of shark teeth being found with plesiosaurimorph specimens.Shimanda K. 2010. A remarkable case of a shark-bitten elasmosaurid plesiosaur. ''Vertebrate Paleontology'', ''30'', 2, 592-597 However it is not certain if the specimens were alive when attacked or whether sharks were feeding on carcasses of deceased plesiosaurimorphs. The shredding teeth of

lamniform The Lamniformes (, from Greek ''lamna'' "fish of prey") are an order of sharks commonly known as mackerel sharks (which may also refer specifically to the family Lamnidae). It includes some of the most familiar species of sharks, such as the g ...

sharks indicates that perhaps they were scavengers or

detrivores Detritivores (also known as detrivores, detritophages, detritus feeders or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...

and that they only fed upon dead plesiosaurs. ''Muraenosaurus'' was an important piece of the ecosystem both as a carnivore and as a source of food to other species.

Gastroliths and Buoyancy

Gastroliths have been a common find among the stomach contents of extinct marine reptiles. Their occurrence has led to two main hypotheses regarding the significance of the rocks. The first proposed usage, as described above, was to crush hard shelled food engulfed by the animal. The second hypothesis is that gastroliths were swallowed in order to help maintain controllable

buoyancy Buoyancy (), or upthrust, is the force exerted by a fluid opposing the weight of a partially or fully immersed object (which may be also be a parcel of fluid). In a column of fluid, pressure increases with depth as a result of the weight of t ...

within the water column. Modeling of ''Cryptoclidus'' body types has indicated that the use of gastroliths as

ballast Ballast is dense material used as a weight to provide stability to a vehicle or structure. Ballast, other than cargo, may be placed in a vehicle, often a ship or the gondola of a balloon or airship, to provide stability. A compartment within ...

is unrealistic.Henderson DM. 2006. Floating point: a computational study of buoyancy, equilibrium, and gastroliths in plesiosaurs. ''Lethaia'', ''39'', 3, 227-244. In order to effect the animal's buoyancy, a Muraenosaur would have to consume over 10% of its body mass in stones. Observed masses of stone collected from plesiosaur stomachs are far lower than the modeled mass required to effect buoyancy. However, from the modeling a new possibility emerged and that is that gastroliths may have helped prevent rolling in animals like ''Muraenosaurus''. Models have indicated that not only do stones reduce pivoting at depth but it is also possible they dampened the ossilations in a plesiosaur's neck, helping provide stability to counter underwater currents.

References

{{Taxonbar, from=Q1179905 Middle Jurassic plesiosaurs of Europe Cryptoclididae Fossil taxa described in 1874 Taxa named by Harry Seeley Oxford Clay Sauropterygian genera