Australochelys
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''Australochelys'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of rhaptochelydian
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 ...
. It is known from one species, ''A. africanus'', that came from the
Elliot Formation The Elliot Formation is a geological formation and forms part of the Stormberg Group, the uppermost geological group that comprises the greater Karoo Supergroup. Outcrops of the Elliot Formation have been found in the northern Eastern Cape, so ...
of
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. The holotype of ''Australochelys'' consists of only a
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
and a fragment of the
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
, which shows both primitive and derived features. Like ''
Proganochelys ''Proganochelys'' is a genus of extinct, primitive stem-turtle. ''Proganochelys'' was named by Georg Baur in 1887 as the oldest turtle in existence at the time. The name ''Proganochelys'' comes from the Greek language, Greek word ''ganos'' meani ...
'', ''Australochelys'' has large orbits and a ventral basioccipital tubercle, but like derived turtles such as casichelydians, a group containing
Cryptodira The Cryptodira (') are a suborder of Testudines that includes most living tortoises and turtles. Cryptodira is commonly called the "Hidden-Neck Turtles" or the "Inside-Neck Turtles". Cryptodira differ from Pleurodira (side-necked turtles) in ...
and
Pleurodira The Pleurodira are one of the two living suborders of turtles, the other being the Cryptodira. The division between these two suborders represents a very deep evolutionary divide between two very different types of turtles. The physical differ ...
, it possesses a sutured basipterygoidal attachment, and a middle ear region partially enclosed laterally. These characteristics show that ''Australochelys'' is more closely related to casichelydians than to ''Proganochelys'', and together with the former, it makes up Rhaptochelydia. The skull of ''Australochelys'' shows that an advanced hearing mechanism of turtles evolved before the appearance of modern turtles.


Etymology

''Australochelys africanus'' was named in 1994 by Eugene S. Gaffney and James W. Kitching. The generic name comes from the
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 ...
''australos'', meaning "south" and ''chelys'', "turtle". The specific name is derived from the Greek ''africanus'', meaning "from Africa".


Description


Skull

The holotype of ''Australochelys'' includes a
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
and a fragment of the
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...
. The skull shares features with both the primitive turtle ''
Proganochelys ''Proganochelys'' is a genus of extinct, primitive stem-turtle. ''Proganochelys'' was named by Georg Baur in 1887 as the oldest turtle in existence at the time. The name ''Proganochelys'' comes from the Greek language, Greek word ''ganos'' meani ...
'' and Casichelydia, a group of derived turtles. The skull shows an advanced hearing mechanism, a feature thought to have evolved with modern turtles.


Distinguishing characteristics

Many features were found among ''Australochelys'' that function to distinguish it from other turtles. Below are the characteristics found by Gaffney and Kitching in 1995 that are unique among all turtles: * an orbit larger in relative size than in any other turtle; * a lacrimal foramen at least three times larger than in ''Proganochelys''; * the external
nare Nare may refer to: People * Daniel Nare, Botswanan football manager * Tadu Teshome Nare (born 2001), Ethiopian runner * Théophile Naré (born 1966), Catholic bishop of Kaya, Burkina Faso Other meanings * Nareh (given name) (or Nare, Nara), Armeni ...
s elongated in contrast to all turtles; * and
vomer The vomer (; ) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxillary bones. The vomer forms ...
s arched dorsally, narrow posteriorly, and very broad anteriorly in a unique configuration. Below are features that are only found in turtles between ''Australochelys'' and ''Proganochelys'', but also present in tetrapods: * the presence of nares formed by the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
divided external; * large interpterygoid vacuity like in ''Proganochelys''; * an unenclosed middle ear region; * a recessed, funnel-shaped, cavum tympani absent; * lacrimal foramen present; * recessus scalae tympani and fenestra perilymphatica absent; * foramen jugulare posterius with at least a medial edge absent; * and the presence of a cultriform process. Below are characteristics found in ''Australochelys'' and Casichelydia but not in ''Proganochelys'': * a sutured basipterygoid articulation; * a stapes that probably does not articulate directly with quadrate but may have attached to a tympanic membrane supported by the acute posterior edge of the quadrate; * a distal end of opisthotic covered by quadrate; * a well-defined canalis cavernosus canal; * few or no palatal teeth; * a distinct crista supraoccipitalis vertical bone sheet; * and a temporal roof that extends posteriorly over opisthotic.


Classification

''Australochelys'' shows a strange mix of basal and derived characteristics. This led Gaffney and Kitching to classify it in two new groups, Rhaptochelydia and Australochelyidae. Rhaptochelydia includes Australochelyidae and Casichelydia, a group including the derived groups
Cryptodira The Cryptodira (') are a suborder of Testudines that includes most living tortoises and turtles. Cryptodira is commonly called the "Hidden-Neck Turtles" or the "Inside-Neck Turtles". Cryptodira differ from Pleurodira (side-necked turtles) in ...
and
Pleurodira The Pleurodira are one of the two living suborders of turtles, the other being the Cryptodira. The division between these two suborders represents a very deep evolutionary divide between two very different types of turtles. The physical differ ...
. Below is a simple
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 ...
found by Gaffney and Kitching in 1995 demonstrating the relationships of ''Australochelys'':


Paleoecology

''Australochelys'' is from the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassicâ ...
Elliot Formation The Elliot Formation is a geological formation and forms part of the Stormberg Group, the uppermost geological group that comprises the greater Karoo Supergroup. Outcrops of the Elliot Formation have been found in the northern Eastern Cape, so ...
, a formation that dates around 190 to 180 million years ago. This makes it the oldest
africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
n turtle, a record that was previously held by a fragmentary specimen from the Gokwe Formation, which was dated to the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
. Other than the fragmentary possible remains, the oldest definite specimens date to the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
of Africa, which shows about a 60 million year gap between themselves and ''Australochelys''. The holotype of ''Australochelys'' was found in a '' Tritylodon'' horizon, in the middle section of the formation.


References

{{Taxonbar, from=Q9162394 Jurassic reptiles of Africa Prehistoric turtle genera Taxa named by Eugene S. Gaffney Taxa named by James Kitching Testudinata Fossil taxa described in 1994