Fraxinisaura
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''Fraxinisaura'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
lepidosauromorph Lepidosauromorpha (in PhyloCode known as ''Pan-Lepidosauria'') is a group of reptiles comprising all diapsids closer to lizards than to archosaurs (which include crocodiles and birds). The only living sub-group is the Lepidosauria, which contains ...
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 ( ...
known from the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
of
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. The only known species is ''Fraxinisaura rozynekae''. It possessed an elongated snout, unique features of the teeth, and an ilium (upper hip bone) which was intermediate in orientation between sphenodontians (the branch of
lepidosaurs The Lepidosauria (, from Greek meaning ''scaled lizards'') is a subclass or superorder of reptiles, containing the orders Squamata and Rhynchocephalia. Squamata includes snakes, lizards, and amphisbaenians. Squamata contains over 9,000 species ...
including the
tuatara Tuatara (''Sphenodon punctatus'') are reptiles endemic to New Zealand. Despite their close resemblance to lizards, they are part of a distinct lineage, the order Rhynchocephalia. The name ''tuatara'' is derived from the Māori language and m ...
) and squamates (the branch including
lizards Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia althou ...
). Based on characteristics of the maxilla, it is considered a close relative of '' Marmoretta'' from the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
of the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...
, resolving a
ghost lineage A ghost lineage is a hypothesized ancestor in a species lineage that has left no fossil evidence yet can be inferred to exist because of gaps in the fossil record or genomic evidence. The process of determining a ghost lineage relies on fossilized ...
between that genus and other Triassic basal lepidosauromorphs.


Discovery

Remains of ''Fraxinisaura'' have only been recovered from the Schumann limestone quarry near Vellberg, Germany. Stratigraphically, it hails from the Untere Graue Mergel Layer of the
Erfurt Formation The Erfurt Formation, also known as the Lower Keuper (German: ''Untere Keuper'', ''Lettenkeuper'', ''Lettenkohle'' or ''Lettenkohlenkeuper''), is a stratigraphic formation of the Keuper group and the Germanic Trias supergroup. It was deposited du ...
, also known as the Lower Keuper. The Lower Keuper preserved a lakeshore environment of
Ladinian The Ladinian is a stage and age in the Middle Triassic series or epoch. It spans the time between Ma and ~237 Ma (million years ago). The Ladinian was preceded by the Anisian and succeeded by the Carnian (part of the Upper or Late Triassic). ...
(late
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma and ...
) age. The holotype of ''Fraxinisaura'' is SMNS 91547, a disarticulated partial skeleton encompassing portions of the skull, vertebrae, hip and limbs. Numerous other isolated bones have also been referred to the taxon. ''Fraxinisaura'' combines the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
words for lizard (''saurus'') and ash tree (''
fraxinus ''Fraxinus'' (), common name, commonly called ash, is a genus of flowering plants in the olive and lilac family, Oleaceae. It contains 45–65 species of usually medium to large trees, mostly deciduous, though a number of Subtropics, subtropic ...
''). This references the nearby village of Eschenau, which means "meadow of ash trees". The species name honors Brigette Rozynek, a fossil collector who donated many Lower Keuper fossils to local museums. ''Fraxinisaura'' was described by Rainer R. Schoch and
Hans-Dieter Sues Hans-Dieter Sues (born January 13, 1956) is a German-born American paleontologist who is Senior Scientist and Curator of Vertebrate Paleontology at the National Museum of Natural History of the Smithsonian Institution in Washington, DC. He receiv ...
in 2018.


Description

The skull of adults was probably 2-2.5 centimeters (.8-1 inches) in length, with an low and elongated snout. Some bones are faintly textured similar to those of kuehneosaurids. The
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
has a short facial process (upwards branch) and a uniquely long premaxillary process (front branch), conditions also known to a lesser extent in ''Marmoretta''. The 22+ maxillary teeth are characteristic as well, being conical, closely spaced, and bearing longitudinal striations at their tips. The teeth are slightly
heterodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. In vertebrates, heterodont pertains to animals where teeth are differentiated into different forms. For example, ...
, with those at the front of the maxilla having more recurved tips than those at the back of the maxilla, or the four at 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 mammal has b ...
. The left and right
frontal Front may refer to: Arts, entertainment, and media Films * ''The Front'' (1943 film), a 1943 Soviet drama film * ''The Front'', 1976 film Music *The Front (band), an American rock band signed to Columbia Records and active in the 1980s and ea ...
and parietal bones at the top of the skull are more robust, flatter, and unfused to their counterparts in contrast to those of ''Marmoretta''. There may have been a small gap where the frontals and parietals meet. The
nasal Nasal is an adjective referring to the nose, part of human or animal anatomy. It may also be shorthand for the following uses in combination: * With reference to the human nose: ** Nasal administration, a method of pharmaceutical drug delivery ** ...
bones at the top of the snout are also longer (from front-to-back) and thinner (from side-to-side) than those of ''Marmoretta''. As with most early lepidosauromorphs, the posterior process (rear branch) of the jugal was very short, leaving the lower temporal fenestra open from below. Preserved postemporal bones, such as the postorbital and squamosal, were smaller and more conservative in shape than those of other early lepidosauromorphs. The rear of the
parasphenoid The parasphenoid is a bone which can be found in the cranium of many vertebrates. It is an unpaired dermal bone which lies at the midline of the roof of the mouth. In many reptiles (including birds), it fuses to the endochondral (cartilage-derived) ...
had a small patch of teeth, a
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...
feature not present in most saurians. There are also three longitudinal rows of small teeth on each pterygoid bone, as well as a transverse row (which is not present in ''Marmoretta''). The ectopterygoid resembles that of ''Sphenodon'' (the modern
tuatara Tuatara (''Sphenodon punctatus'') are reptiles endemic to New Zealand. Despite their close resemblance to lizards, they are part of a distinct lineage, the order Rhynchocephalia. The name ''tuatara'' is derived from the Māori language and m ...
). The dentary has a small expansion at the tip, forming a "chin" similar to that of sphenodontians. The elongated, closely spaced dentary teeth are similar to those of the maxilla. Dentition is pleurodont, like other basal lepidosauromorphs and lizards, but unlike sphenodontians.
Vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
are similar to those of ''Marmoretta'' and kuehneosaurs, being amphicoelous (concave at both ends) and lacking a hole for the
notochord In anatomy, the notochord is a flexible rod which is similar in structure to the stiffer cartilage. If a species has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a chordate. The notochord consis ...
. 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 coracoid are fused into a scapulocoracoid, while the interclavicle was large, arrow-shaped, and heavily textured. Like other lepidosauromorphs, the
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
is twisted, constricted in the middle, and possesses an
entepicondylar foramen The entepicondylar foramen is an opening in the distal (far) end of the humerus (upper arm bone) present in some mammals. It is often present in primitive placentals, such as the enigmatic Madagascan ''Plesiorycteropus''. In most Neotominae and al ...
but not an ectepicondylar one. The ilium has a distinctively long, leaf-shaped iliac blade which projects up and back at a 45 degree angle, intermediate between the horizontal iliac blade of
lizards Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia althou ...
, and the vertical iliac blade of sphenodontians. ''
Sophineta ''Sophineta'' is an extinct genus of basal lepidosauromorph reptile known from the Early Triassic (late Olenekian age) of Małopolska Province, southern Poland. It contains a single species, ''Sophineta cracoviensis''. Discovery ''Sophineta'' ...
'' and ''
Gephyrosaurus ''Gephyrosaurus'' is a genus of early rhynchocephalian first described and named in 1980 by Susan E. Evans. They are distantly related to the extant ''Sphenodon'' (tuatara of New Zealand) with which they shared a number of skeletal features incl ...
'' have a similarly shaped intermediate ilium. Recovered hindlimb material generally resembles that of generalized
lepidosaurs The Lepidosauria (, from Greek meaning ''scaled lizards'') is a subclass or superorder of reptiles, containing the orders Squamata and Rhynchocephalia. Squamata includes snakes, lizards, and amphisbaenians. Squamata contains over 9,000 species ...
like ''Sphenodon'', though the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
is somewhat more robust.


Classification

''Fraxinisaura'' does not preserve a
quadrate Quadrate may refer to: * Quadrate bone * Quadrate (heraldry) * Quadrate lobe of liver * Quadrate tubercle The quadrate tubercle is a small tubercle found upon the upper part of the femur. It serves as a point of insertion of the quadratus femori ...
, sternum, or metatarsal V, bones which are useful for determining whether a reptile is a lepidosauromorph or not. However, it does possess an entepicondylar foramen of the humerus as well as pleurodont dentition, both of which are characteristic lepidosauromorph features. The shape of the maxilla also links it to ''Marmoretta'', helping to fill the small
ghost lineage A ghost lineage is a hypothesized ancestor in a species lineage that has left no fossil evidence yet can be inferred to exist because of gaps in the fossil record or genomic evidence. The process of determining a ghost lineage relies on fossilized ...
which lies between ''Marmoretta'' (from the mid to late
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
) and other basal lepidosauromorphs (which appeared in 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 ...
). This classification scheme was supported by a phylogenetic analysis using the data matrix of Ezcurra ''et al.'' (2014). The ''Fraxinisaura'' + ''Marmoretta'' clade was tentatively considered to be closer to ''Sophineta'' and rhynchocephalians than to lizards, but lepidosauromorph relations were largely reduced into a polytomy upon the addition of ''
Kuehneosaurus ''Kuehneosaurus'' is an extinct genus of Late Triassic kuehneosaurid reptile known from the Late Triassic ( Norian stage) of the Penarth Group of southwest England and the Steinmergel Group of Luxembourg. Temperature at this stage and region w ...
'' into the analysis. Adding ''Fraxinisaura'' to another lepidosauromorph analysis (
Evans Evans may refer to: People *Evans (surname) *List of people with surname Evans Places United States *Evans Island, an island of Alaska *Evans, Colorado *Evans, Georgia *Evans County, Georgia *Evans, New York *Evans Mills, New York *Evans City, ...
and Borsuk-Białynicka (2009)) also led to a large polytomy. Below is a portion of the cladogram produced by adding ''Fraxinisaura'' (but not ''Kuehneosaurus'') to the analysis of Ezcurra ''et al.'' (2014):


References

{{Taxonbar, from1=Q56254434, from2=Q56254445 Middle Triassic reptiles of Europe Triassic lepidosauromorphs Fossil taxa described in 2018 Prehistoric reptile genera