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''Mesosaurus'' (meaning "middle lizard") is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
of reptile from the
Early Permian 01 or '01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * 01'' (Richard Müller album), 2001 * ''01'' (Son of Dave album), 2000 * ''01'' (Urban Zakapa album), 2011 * ''O1'' (Hiroyuki Sawano ...
of southern
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the souther ...
. Along with it, the genera ''
Brazilosaurus ''Brazilosaurus'' is an extinct genus of mesosaur which lived during the early Permian ( Artinskian stage) of what is now Brazil. It is known from specimen BSPG 1965 I 131, a single skeleton recovered from the Assistencia Member of the Irati Form ...
'' and '' Stereosternum'', it is a member of the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Mesosauridae and the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
Mesosauria. ''Mesosaurus'' was long thought to have been one of the first
marine reptile Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. The earliest marine reptile mesosaurus (not to be confused with mosasaurus), arose in the Permian period during ...
s, although new data suggests that at least those of Uruguay inhabited a hypersaline water body, rather than a typical marine environment. In any case, it had many adaptations to a fully aquatic lifestyle. It is usually considered to have been anapsid, although Friedrich von Huene considered it to be a
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes rep ...
, and this hypothesis has been revived recently.


Discovery and naming

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
of ''M. tenuidens'', MNHN 1865-77, is nicknamed the " Griqua ''Mesosaurus''" and it was found in a Griqua hut in
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring count ...
, likely in Kimberley, Northern Cape around 1830 and was being used as a pot lid. The circumstances of its discovery and how it was taken from its previous owners in South Africa are unknown, but what is known is that the specimen eventually surfaced in the collection of the
French French (french: français(e), link=no) may refer to: * Something of, from, or related to France ** French language, which originated in France, and its various dialects and accents ** French people, a nation and ethnic group identified with Franc ...
palaeontologist Paul Gervais during the 1860s and he designated it as the holotype of a new genus and species he named ''Mesosaurus tenuidens'' in 1865. Since then, ''Mesosaurus'' remains have also been identified from
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the souther ...
and were first identified in 1908 as belonging to a second species, ''M. brasiliensis'', by J. H. MacGregor. Later studies have shown that ''M. brasiliensis'' was the same animal as ''M. tenuidens'', which remains as the single valid species of ''Mesosaurus to this day. Two other species of mesosaurids have since been described, which are '' Stereosternum''Cope, E.D. (1885). A contribution to the vertebrate paleontology of Brazil. ''Proceedings of the American Philosophical Society'' 25, 7-15. and ''
Brazilosaurus ''Brazilosaurus'' is an extinct genus of mesosaur which lived during the early Permian ( Artinskian stage) of what is now Brazil. It is known from specimen BSPG 1965 I 131, a single skeleton recovered from the Assistencia Member of the Irati Form ...
'', which are also considered to be synonyms of ''Mesosaurus tenuidens'' according to Piñeiro ''et al.'' (2021).


Description

''Mesosaurus'' had a long skull that was larger than that of ''Stereosternum'' and had longer teeth. The teeth are angled outwards, especially those at the tips of the jaws. The bones of the postcranial skeleton are thick, having undergone pachyostosis. ''Mesosaurus'' is unusual among reptiles in that it possesses a
cleithrum The cleithrum (plural cleithra) is a membrane bone which first appears as part of the skeleton in primitive bony fish, where it runs vertically along the scapula. Its name is derived from Greek κλειθρον = "key (lock)", by analogy with " cl ...
. A cleithrum is a type of dermal bone that overlies the
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
, and is usually found in more primitive bony fish and tetrapods. The head of the interclavicle of ''Mesosaurus'' is triangular, unlike those of other early reptiles, which are diamond-shaped.


Palaeobiology

''Mesosaurus'' was one of the first reptiles known to have returned to the water after early
tetrapod Tetrapods (; ) are four-limb (anatomy), limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant taxon, extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (p ...
s came to land in the Late Devonian or later in the Paleozoic. It was around in length, with webbed feet, a streamlined body, and a long tail that may have supported a fin. It probably propelled itself through the water with its long hind legs and flexible tail. Its body was also flexible and could easily move sideways, but it had heavily thickened ribs, which would have prevented it from twisting its body. ''Mesosaurus'' had a small
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, t ...
with long jaws. The
nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called t ...
s were located at the top, allowing the creature to breath with only the upper side of its head breaking the surface, in a similar manner to a modern
crocodile Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant ...
. The teeth were originally thought to have been straining devices for the filter feeding of
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a cr ...
ic organisms. However, this idea was based on the assumption that the teeth of ''Mesosaurus'' were numerous and close together in the jaws. Newly examined remains of ''Mesosaurus'' show that it had fewer teeth and that the dentition was suitable for catching small nektonic prey such as crustaceans. The pachyostosis seen in the bones of ''Mesosaurus'' may have enabled it to reach neutral buoyancy in the upper few meters of the water column. The additional weight may have stabilized the animal at the water's surface. Alternatively, it could have given ''Mesosaurus'' greater momentum when gliding underwater. While many features suggest a wholly aquatic lifestyle, ''Mesosaurus'' may have been able to move onto land for short periods of time. Its elbows and ankles were restricted in their movement, making walking appear impossible. It is more likely that if ''Mesosaurus'' moved onto land, it would push itself forward in a similar way to living female
sea turtle Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerhe ...
s when nesting on beaches. Clearly amniote-type fossil
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...
s of ''Mesosaurus'' in an advanced stage of development (i.e.
fetus A fetus or foetus (; plural fetuses, feti, foetuses, or foeti) is the unborn offspring that develops from an animal embryo. Following embryonic development the fetal stage of development takes place. In human prenatal development, fetal develo ...
es) have been discovered in Uruguay and Brazil. These fossils are the earliest record of amniote fetuses, although amniotes are inferred to have had their typical reproductive strategy since their first appearance in the
Late Carboniferous Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effe ...
. Prior to their description, the oldest known amniote fetuses were from the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
. One isolated coiled fetus called FC-DPV 2504 is not surrounded by
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an a ...
eggshells, suggesting that the glands in the
oviduct The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, ...
of ''Mesosaurus'' and probably all Paleozoic amniotes were not able to secrete calcium carbonate, in contrast to post-paleozoic
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avi ...
s. This would explain the scarcity of egg fossils in the paleozoic amniote fossil record. One ''Mesosaurus'' specimen called MCN-PV 2214 comprises a medium-size adult with a small individual in its rib cage which is interpreted as a fetus ‘ in utero’, even suggesting that ''Mesosaurus'' like many other
marine reptile Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. The earliest marine reptile mesosaurus (not to be confused with mosasaurus), arose in the Permian period during ...
s, gave live birth. If this interpretation is correct, this specimen would represent the earliest known example of
viviparity Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
in the fossil record. The isolated fetus FC-DPV 2504, however, rather points to an
ovoviviparous Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop insi ...
reproduction strategy in ''Mesosaurus''. A study on vertebral column proportions suggested that, while young ''Mesosaurus'' might have been fully aquatic, adult animals spent some time on land. This is supported by the rarity of adult animals in aquatic settings, and a coprolite possessing drying fractures. However, how terrestrial these animals were is difficult to say, as their pachyostosis and other adaptations for an aquatic lifestyle would have made foraging on land difficult.


Distribution

''Mesosaurus'' was significant in providing evidence for the
theory A theory is a rational type of abstract thinking about a phenomenon, or the results of such thinking. The process of contemplative and rational thinking is often associated with such processes as observational study or research. Theories may ...
of
continental drift Continental drift is the hypothesis that the Earth's continents have moved over geologic time relative to each other, thus appearing to have "drifted" across the ocean bed. The idea of continental drift has been subsumed into the science of pla ...
, because its remains were found in southern Africa,
Whitehill Formation The Whitehill Formation, alternatively written as White Hill Formation and formerly known as White Band or Whitehill or White Hill Member, is a regional Early Permian (Artinskian to Kungurian, dating to around 282 to 275 Ma) geologic formation b ...
, and eastern South America ( Melo Formation, Uruguay and Irati Formation, Brazil), two widely separated regions. As ''Mesosaurus'' was a coastal animal, and therefore less likely to have crossed the
Atlantic Ocean The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth#Surface, Earth's surface and about 29% of its water surface area. It is known to separate the "Old World" of Afr ...
, this distribution indicated that the two continents used to be joined together.


Gallery

Mesosaurus tenuidens 1.jpg, Fossil from
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
Mesosaurus.jpg, Fossil in Milan Dual Mesosaurus.jpg, Fossil in Louisiana Mesosaurus BW.jpg, Restoration Mesosaurus.png, Early reconstruction of the skeleton of ''M. brasiliensis'' showing many small teeth in the jaws (MacGregor, 1908).MacGregor, J.H. (1908) ''Mesosaurus brasiliensis'' nov. sp. IN: White, I.C. (1908) Commission for Studies on Brazilian Coal Mines - Final Report; (Bilingual report, Portuguese & English), Imprensa Nacional, Rio de Janeiro, Brazil, 617 p.: Part II, pp. 301-336. Mesosaurus fetus Pineiro et al (2016) PeerJ 4-e2036 fig S1.png, Fossil of unhatched juvenile or fetus of ''Mesosaurus tenuidens'' (FC-DPV 2504) from Uruguay Mesosaurus Fossil bei Keetmanshoop, Namibia.jpg, Skeleton molds in whitish weathering shales of the Whitehill Formation, Keetmanshoop,
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and e ...
Wegener fossils-mapped.png, Distribution of four Permian and Triassic fossil groups used as biogeographic evidence for continental drift, and land bridging. Location of ''Mesosaurus'' remains shown by green squares


References


Further reading

* Parker, Steve. Dinosaurus: the complete guide to dinosaurs. Firefly Books Inc, 2003. Pg. 90 * * * * {{Taxonbar, from=Q132822 Prehistoric marine reptiles Prehistoric reptile genera Permian reptiles of Africa Fossils of Namibia Fossils of South Africa Permian reptiles of South America Permian Brazil Fossils of Brazil Permian Uruguay Fossils of Uruguay Paraná Basin Fossil taxa described in 1865 Taxa named by Paul Gervais