Saurosphargid
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Saurosphargidae is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
marine reptile Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. Only about 100 of the 12,000 extant reptile species and subspecies are classed as marine reptiles, including mari ...
s known from the
Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between 251.9 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which ...
(
Olenekian In the geologic timescale, the Olenekian is an age (geology), age in the Early Triassic epoch (geology), epoch; in chronostratigraphy, it is a stage (stratigraphy), stage in the Lower Triassic series (stratigraphy), series. It spans the time betw ...
stage) and early
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epoch (geology), epochs of the Triassic period (geology), period or the middle of three series (stratigraphy), series in which the Triassic system (stratigraphy), system is di ...
(
Anisian In the geologic timescale, the Anisian is the lower stage (stratigraphy), stage or earliest geologic age, age of the Middle Triassic series (stratigraphy), series or geologic epoch, epoch and lasted from million years ago until million years ag ...
stage) of
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
and
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. The
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
of the family is '' Saurosphargis'', named by
Friedrich von Huene Baron Friedrich Richard von Hoyningen-Huene (22 March 1875 – 4 April 1969) was a German nobleman paleontologist who described a large number of dinosaurs, more than anyone else in 20th-century Europe. He studied a range of Permo-Carbonife ...
in 1936 based on a single specimen collected from the lower Anisian Lower Muschelkalk of Gogolin,
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...
-- a section of 12 incomplete
back 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 vertebrae of intermediate size between the cervical and lumbar ve ...
with
rib In vertebrate anatomy, ribs () are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the thoracic cavity, enabling the lungs to expand and thus facilitate breathing by expanding the ...
s. The generic name of ''Saurosphargis'' is derived from
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
''sauros'', "lizard," and ''sphargis'', the old genus name for the leatherback turtle, in reference to the dorsal
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amph ...
"body armor" and broadened ribs forming a closed chest rib basket, traits that are seemingly transitional between
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s and other reptiles. However, due to the only known specimen' destruction during
World War II World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...
, many authors considered ''Saurosphargis'' to be a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
''. Nevertheless, even based on surviving descriptions alone, many differences were noted even from its closest known relative at the time, '' Eusaurosphargis dalsassoi'' from the Anisian-Ladinian boundary of the southern
Alps The Alps () are some of the highest and most extensive mountain ranges in Europe, stretching approximately across eight Alpine countries (from west to east): Monaco, France, Switzerland, Italy, Liechtenstein, Germany, Austria and Slovenia. ...
. Various hypotheses existed for the affinities of these species, and together with ''
Helveticosaurus ''Helveticosaurus'' is an extinct genus of diapsid marine reptile known from the Middle Triassic (Anisian-Ladinian boundary) of southern Switzerland. It contains a single species, ''Helveticosaurus zollingeri'', known from the nearly complete h ...
'', they were originally thought to be
placodont Placodonts (" tablet teeth") are an extinct order of marine reptiles that lived during the Triassic period, becoming extinct at the end of the period. They were part of Sauropterygia, the group that includes plesiosaurs. Placodonts were generall ...
s, but later studies suggested relatedness to other
sauropterygia Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
ns and / or
ichthyopterygia Ichthyopterygia ("fish flippers") was a designation introduced by Richard Owen, Sir Richard Owen in 1840 to designate the Jurassic ichthyosaurs that were known at the time, but the term is now used more often for both true Ichthyosauria and their ...
ns. It wasn't until nearly a century after the discovery of ''Saurosphargis'', that other specimens closely related to it were found in
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. In 2011, Li ''et al.'' described and named three of these specimens, '' Sinosaurosphargis yunguiensis'', referring to this species being the Chinese version of ''Saurosphargis''. The specimens of ''Sinosaurosphargis'' were collected from Member II of the
Guanling Formation The Guanling Formation is a Middle Triassic (Anisian or Pelsonian in the regional chronostratigraphy) geologic Formation (geology), formation in southwestern China. Geology The formation encompasses two members. The first member is primarily c ...
, dating to the Pelsonian (mid-late
Anisian In the geologic timescale, the Anisian is the lower stage (stratigraphy), stage or earliest geologic age, age of the Middle Triassic series (stratigraphy), series or geologic epoch, epoch and lasted from million years ago until million years ag ...
). Based on very detailed descriptions and figures of ''Saurosphargis'' in Huene (1936), and based on comparisons with the well-preserved ''Sinosauropshargis'', Li ''et al.'' (2011) resurrected ''Saurosphargis volzi'' as a valid species, and erected the family Saurosphargidae to contain the two. Furthermore, they reported the discovery of material referable to ''Saurosphargis'' from the Lower Muschelkalk of
Winterswijk Winterswijk (; also known as ''Winterswiek'' or ''Wenters'') is a Municipalities of the Netherlands, municipality and a town in the eastern Netherlands. It has a population of and is situated in the Achterhoek, which lies in the easternmost part ...
, the
Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
, that is under preparation. ''Saurosphargis'' and ''Sinosaurosphargis'' were included in a
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
, and were recovered in a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
Saurosphargidae which was found to be the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of
Thalattosauriformes Thalattosauria (Greek for "sea lizards") is an extinct order of marine reptiles that lived during the Triassic Period. Thalattosaurs were diverse in size and shape, and are divided into two superfamilies: Askeptosauroidea and Thalattosauroidea. ...
.
Sauropterygia Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
was recovered as the sister taxon of their combined group, with a clade formed by ''Eusaurosphargis'' and ''Helveticosaurus'' in a position basal to it. In 2014, Li ''et al.'' described a second species of '' Largocephalosaurus'', that was initially interpreted as an
eosauropterygia Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
n due to the limited preparation of the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
of its type species ''L. polycarpon''. The second species ''L. qianensis'', known from three specimens, and the newly prepared portions of ''L. polycarpon''
postcranial The postcranium ("behind the cranium"; plural: postcrania) or postcranial skeleton in zoology and vertebrate paleontology is the skeleton apart from the skull. The postcranium encompasses the axial skeleton, which includes the entirety of the verte ...
skeleton, revealed the same osteoderm "body armor" and broadened rib basket that are unique to saurosphargids. Thus, ''Largocephalosaurus'' was reinterpreted as a saurosphargid and a phylogenetic analysis recovered both species, forming a monophyletic ''Largocephalosaurus'', as basal to the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
formed by ''Saurosphargis'' and ''Sinosaurosphargis''. All four known specimens of ''Largocephalosaurus'' came from the Luoping Biota, from Member II of the Guanling Formation, that yielded ''Sinosaurosphargis'', alongside various invertebrates, fish, basal
ichthyosaur Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...
s, '' Atopodentatus'', and several species of
eosauropterygia Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
n, including both
pachypleurosaur Pachypleurosauria is an extinct clade of primitive sauropterygian reptiles from the Triassic period. Pachypleurosaurs vaguely resembled aquatic lizards, with elongate forms ranging in size from , with small heads, long necks, paddle-like limbs, a ...
s and
nothosaurid Nothosauridae are an extinct family of carnivorous aquatic sauropterygian reptiles from the Triassic time period of China, France, Germany, Israel, Italy, Netherlands, Russia, Switzerland, and northern Africa. Phylogeny The cladogram shown bel ...
s.


Phylogeny

The following
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
is simplified after Li ''et al.'' (2014)
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
, showing interrelationships of all known Saurosphargidae species, and the placement of the clade within
Sauria Sauria is the clade of diapsids containing the most recent common ancestor of Archosauria (which includes crocodilians and birds) and Lepidosauria (which includes squamates and the tuatara), and all its descendants. Since most molecular phyl ...
. ''Saurosphargis'' was coded solely based on the holotype. The removal / inclusion of
Ichthyopterygia Ichthyopterygia ("fish flippers") was a designation introduced by Richard Owen, Sir Richard Owen in 1840 to designate the Jurassic ichthyosaurs that were known at the time, but the term is now used more often for both true Ichthyosauria and their ...
was found to affect the
topology Topology (from the Greek language, Greek words , and ) is the branch of mathematics concerned with the properties of a Mathematical object, geometric object that are preserved under Continuous function, continuous Deformation theory, deformat ...
the most - switching the positions of the ''Eusaurosphargis''+''Helveticosaurus'' and
Thalattosauriformes Thalattosauria (Greek for "sea lizards") is an extinct order of marine reptiles that lived during the Triassic Period. Thalattosaurs were diverse in size and shape, and are divided into two superfamilies: Askeptosauroidea and Thalattosauroidea. ...
clades, and altering the positions of several taxa within
Eosauropterygia Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
, which are not shown. The Saurosphargiformes are sometimes grouped together with Sauropterygia in a clade called Sauropterygomorpha. Wolniewicz ''et al.'' (2023) recovered saurosphargids nested within Sauropterygia, more closely related to
eosauropterygia Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic diapsid reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosau ...
ns than to placodonts:


References

{{Taxonbar, from=Q20984193 Middle Triassic reptiles Fossils of China Middle Triassic first appearances Middle Triassic extinctions Prehistoric reptile families Sauria