Leonerasaurus
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''Leonerasaurus'' is a basal
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
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lon ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
. Currently, there is only one
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
known, named ''L. taquetrensis'' by Diego Pol, Alberto Garrido and Ignacio A. Cerda in
2011 File:2011 Events Collage.png, From top left, clockwise: a protester partaking in Occupy Wall Street heralds the beginning of the Occupy movement; protests against Libyan dictator Muammar Gaddafi, who was killed that October; a young man celebrate ...
. The
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
, an incomplete
subadult A juvenile is an individual organism that has not yet reached its adult form, sexual maturity or size. Juveniles can look very different from the adult form, particularly in colour, and may not fill the same niche as the adult form. In many org ...
individual, was found in the Las Leoneras Formation in
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. This formation is probably
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 Triassic-J ...
in age. ''Leonerasaurus'' was a small non-
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
sauropodomorph, showing an unusual combination of basal and derived characters. This indicates that the evolution of early sauropodomorphs witnessed a great degree of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
.


Discovery

The fossils assigned to ''Leonerasaurus'' were found near Cañadón Las Leoneras (an affluent of the left margin of the Chubut river), southeast of Sierra de Taquetrén,
Chubut Province Chubut ( es, Provincia del Chubut, ; cy, Talaith Chubut) is a province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa Cruz Province), the Andes ra ...
, Central
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and gl ...
, Argentina. This formation is probably early Jurassic in age, interpreted as
Pliensbachian The Pliensbachian is an age of the geologic timescale and stage in the stratigraphic column. It is part of the Early or Lower Jurassic Epoch or Series and spans the time between 190.8 ± 1.5 Ma and 182.7 ± 1.5 Ma (million years ago). The Plien ...
to
Toarcian The Toarcian is, in the ICS' geologic timescale, an age and stage in the Early or Lower Jurassic. It spans the time between 182.7 Ma (million years ago) and 174.1 Ma. It follows the Pliensbachian and is followed by the Aalenian. The Toarcian ...
or late
Sinemurian In the geologic timescale, the Sinemurian is an age and stage in the Early or Lower Jurassic Epoch or Series. It spans the time between 199.3 ± 2 Ma and 190.8 ± 1.5 Ma (million years ago). The Sinemurian is preceded by the Hettangian and is ...
to Toarcian. The volcanic facies of the overlying Lonco Trapial Formation is certainly 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 ...
, so that the younger boundary of the Las Leoneras Formation is well constrained. However, a
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
affinity cannot be rules out, because the lower constraint of the formation is not well-defined.


Etymology

The generic name is derived from ''Leoneras'', in reference to the lithostratigraphic unit where this taxon was found, and ''saurus'', "lizard" (Latinized Greek). The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
''taquetrensis'' refers to the Sierras de Taquetrén, where Las Leoneras Formation crops out in Central Patagonia.


Description

''Leonerasaurus taquetrensis'' is known from one incomplete individual. Parts of a
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
and some teeth, neck and trunk
vertebra 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 ...
e, a
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
, parts of the
pectoral Pectoral may refer to: * The chest region and anything relating to it. * Pectoral cross, a cross worn on the chest * a decorative, usually jeweled version of a gorget * Pectoral (Ancient Egypt), a type of jewelry worn in ancient Egypt * Pectoralis ...
(shoulder) and
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
(hip) girdle as well as several limb bones were found. Much of the remains were found in articulation. It was only 2.40 meters long, 1 m high and weighing about 70 kg.


Dentary and teeth

Of the skull only the anterior part of the right
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
was found. Near the area where it touches the contralateral element at the tip of the lower jaw (the
symphysis A symphysis (, pl. symphyses) is a fibrocartilaginous fusion between two bones. It is a type of cartilaginous joint, specifically a secondary cartilaginous joint. # A symphysis is an amphiarthrosis, a slightly movable joint. # A growing together ...
) the bone is straight and only gently arched medially, as is seen in basal sauropods. More derived sauropods ( eusauropods) have medially broadly arching symphyseal regione and anterior portions of the tooth row. The ventral (lower) edge of the dentary is damaged, but does not appear to be ventrally deflected at the symphysis as in some basal sauropodomorphs such as
Plateosaurus ''Plateosaurus'' (probably meaning "broad lizard", often mistranslated as "flat lizard") is a genus of plateosaurid dinosaur that lived during the Late Triassic period, around 214 to 204 million years ago, in what is now Central and Northern Eur ...
. Also, a longitudinal ridge that is seen in some basal sauropodomorphs (e.g., ''
Massospondylus ''Massospondylus'' ( ; from Greek, (massōn, "longer") and (spondylos, "vertebra")) is a genus of sauropodomorph dinosaur from the Early Jurassic. (Hettangian to Pliensbachian ages, ca. 200–183 million years ago). It was described by S ...
'', ''
Coloradisaurus ''Coloradisaurus'' (meaning "Los Colorados lizard") is a genus of massospondylid sauropodomorph dinosaur. It lived during the Late Triassic period ( Norian stage) in what is now La Rioja Province, Argentina. It is known from two specimens col ...
'' and ''
Plateosaurus ''Plateosaurus'' (probably meaning "broad lizard", often mistranslated as "flat lizard") is a genus of plateosaurid dinosaur that lived during the Late Triassic period, around 214 to 204 million years ago, in what is now Central and Northern Eur ...
'') is not preserved in ''Leonerasaurus''. However, this may be related to the damaged state of the specimen. On the dentary, 13 teeth or tooth fragments were found. There are two empty
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * ...
, so that the total tooth count on one side was at least 15. Three teeth were found isolated near the dentary. The teeth and alveoli are angled forward (procumbent) by ~60°, similar to eusauropods, but also to juveniles of ''
Mussaurus ''Mussaurus'' (meaning " mouse lizard") is a genus of herbivorous sauropodomorph dinosaur that lived in southern Argentina during the Early Jurassic, with a maximum age of 192.78 ± 0.14 Ma. It receives its name from the small size of the sk ...
''. Tooth height and width decreases from the front to the back, and neighboring teeth overlapping each other. The edges of the teeth in the front of the jaw likely were not serrated, or at least only at the crown tips. This is usually the case in eusauropods. Teeth further back in the dentary are nearly all damaged, but a not erupted tooth has large denticles similar to basal sauropodomorphs. This pattern of teeth without denticles in front, and teeth with denticles in the back of the lower jaw is also seen in juveniles of ''Mussaurus'' and ''
Melanorosaurus ''Melanorosaurus'' (meaning "Black Mountain Lizard", from the Greek ''melas/'', "black", ''oros/'', "mountain" + ''/'', "lizard") is a genus of basal sauropodomorph dinosaur that lived during the Late Triassic period. A herbivore from South Af ...
''. Also, in contrast to eusauropod teeth, there are no high-angle wear facets on the teeth of ''Leonorasaurus''. The anterior teeth are spoon-shaped, with the outer surface convex (bulging out), the inner one concave (hollowed out), again resembling basal sauropods. The surface texture, however, is more similar to basal sauropodomorphs, and not to sauropods.


Vertebrae

The neck of ''Leonerasaurus'' is known almost completely, only the
atlas An atlas is a collection of maps; it is typically a bundle of maps of Earth or of a region of Earth. Atlases have traditionally been bound into book form, but today many atlases are in multimedia formats. In addition to presenting geographic ...
is lost. The nine remaining cervicals were found articulated with the first five complete
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal co ...
(trunk) vertebrae. Additionally, a probable sixth dorsal and an articulated group of three mid-dorsals were found. Fragments of more posterior dorsals and of ribs were also recovered. The sacrum (fused vertebrae that form the core of the hip), consisting of four vertebrae, is nearly complete, but no bones of the tail was recovered. The overall shape of the neck vertebrae is typical for basal sauropodomorphs, but Pol et al. interpret the remains of the neural arches to indicate a more sauropod-like shape. Some characters, however, show an intermediate development. In the trunk, the vertebrae show typical non-sauropod characters, such as relatively long and low neural arches with a narrow anterior ridge (the anteriormost dorsal vertebrae have slightly high neural arches), an anteriorly placed parapophysis (one of the articulation points for the ribs), and lack of or less strong development of certain laminae (thin ridges). The diapophysis (the second articulation for the rib) is also in the same position as in basal sauropodomorphs. A lamina connecting the diapophysis to the
prezygapophysis The articular processes or zygapophyses (Greek ζυγον = "yoke" (because it links two vertebrae) + απο = "away" + φυσις = "process") of a vertebra are projections of the vertebra that serve the purpose of fitting with an adjacent vertebr ...
on all dorsals, but not in no-sauropod sauropodomorphs, is present in the posterior dorsals.


Phylogenetic position

''Leonerasaurus'' appears to be belong into the
Anchisauria Anchisauria is an extinct clade of sauropodomorph dinosaurs that lived from the Late Triassic to the Late Cretaceous. The name Anchisauria was first used Haekel and defined by Galton and Upchurch in the second edition of ''The Dinosauria''. It is ...
, as the closest sister
taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
to the group sauropods (following Yates) + ''Melanorosaurus'' (''M.'' is seen as a sauropod by some researchers). A
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
after Pol, Garrido & Cerda, 2011, illustrates a possible placing of ''Leonerasaurus'' in Sauropodomorpha: The following cladogram shows the position of ''Leonerasaurus'' within Massopoda, according to Oliver W. M. Rauhut and colleagues, 2020:


References

{{Taxonbar, from=Q135102 Anchisauria Dinosaur genera Early Jurassic dinosaurs of South America Jurassic Argentina Fossils of Argentina Cañadón Asfalto Basin Fossil taxa described in 2011