''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 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 ...

plesiosaur The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared ...

reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...

from the Oxford Clay of Southern England. 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 of the middle Jurassic. Charles E. Leeds collected the first ''Muraenosaurus'' which was then described by H. G. Seeley.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 Charles J. Leeds was the 37th mayor of New Orleans (November 30, 1874 – December 19, 1876). Leeds was a member of the white supremacist organization White League and had furnished its members with artillery and small arms during their insurre ...

’ 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 Leedsi''. ''M. leedsi'' 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 ''Picrocleidus beloclis''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. ''Muraenosaurus reedii'' was discovered in Wyoming 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. leedsi'' 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 The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...

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

Picrocleidus ''Picrocleidus'' is an extinct genus of plesiosaur. It is known only from the type species ''P. beloclis'' from the Middle Jurassic Oxford Clay Formation (Callovian stage) of the United Kingdom.Andrews, Charles W 1910. A descriptive catalogue of ...

'' 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 (singular: 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 ...

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. 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. The neural spines have elongated anteroposteriorly and compressed vertically. The pectoral girdle in ''Muraenosaurus'' is broader than in most plesiosaurs and helped to situate the animal as a member of ''Cryptoclididae''. The

coracoids A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...

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 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 fat ...

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

Elasmosauridae Elasmosauridae is an extinct family of plesiosaurs, often called elasmosaurs. They had the longest necks of the plesiosaurs and existed from the Hauterivian to the Maastrichtian stages of the Cretaceous, and represented one of the two groups of p ...

instead of

Cryptoclididae Cryptoclididae is a family (biology), 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 ...

. 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 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 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. ''Kimmerosaurus'' is most closely related to ''Tatenectes''. Discovery There are very few fossil remains of ''Kimmerosaurus'' known. ...

'' 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 fore limbs especially resemble the flippers of '' Pliosauridae'', an extinct group of less elongate marine reptiles that shared the seas with plesiosaurs, more than they resemble the typical plesiosaur morphology. ''

Cryptoclidus ''Cryptoclidus'' ( ) is a genus of plesiosaur reptile from the Middle Jurassic Period (geology), period of England, France, and Cuba. Discovery ''Cryptoclidus'' was a plesiosaur whose specimens include adult and juvenile skeletons, and remai ...

'', 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 (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...

arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...

, gastropods and foraminifera 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 fish and

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

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

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 eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evolut ...

. It was proposed that plesiosaurimophs would swim with a straight neck and attack pelagic 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 were dominated by pliosaurs, apex predators 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 gre ...

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 invertebrat ...

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 A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In oth ...

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 within the water column. Modeling of ''Cryptoclidus'' body types has indicated that the use of gastroliths as ballast 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 Cryptoclidids Fossil taxa described in 1874 Taxa named by Harry Seeley Oxford Clay Sauropterygian genera