Stenaulorhynchus stockleyi
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''Stenaulorhynchus'' (possibly meaning "narrow tube beak") 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 living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of hyperodapedontid rhynchosaur 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 an ...
(late
Anisian In the geologic timescale, the Anisian is the lower stage or earliest age of the Middle Triassic series or epoch and lasted from million years ago until million years ago. The Anisian Age succeeds the Olenekian Age (part of the Lower Triassic ...
stage) deposits of
Tanganyika Territory Tanganyika was a colonial territory in East Africa which was administered by the United Kingdom in various guises from 1916 to 1961. It was initially administered under a military occupation regime. From 20 July 1922, it was formalised into a L ...
,
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
. It was found in the Lifua Member of the Manda Formation in the
Karoo Supergroup The Karoo Supergroup is the most widespread stratigraphic unit in Africa south of the Kalahari Desert. The supergroup consists of a sequence of units, mostly of nonmarine origin, deposited between the Late Carboniferous and Early Jurassic, a peri ...
. It was named and first described by Sidney Henry Haughton in
1932 Events January * January 4 – The British authorities in India arrest and intern Mahatma Gandhi and Vallabhbhai Patel. * January 9 – Sakuradamon Incident: Korean nationalist Lee Bong-chang fails in his effort to assassinate Emperor Hir ...
. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specim ...
is ''Stenaulorhynchus stockleyi'', a beaked herbivore measuring 1–6 meters in length.


Description


Dentition

The teeth of ''Stenaulorhynchus'' were conical, pointed, and composed mostly of dentine, although new unworn teeth may have had a thin layer of enamel. They were set deeply into and fused with the jaw bones. They are arranged with two-to-several rows of teeth on top and only a few on bottom. The middle row of maxillary teeth only occupied the posterior third of the jaw while the other rows extended all the way forwards and sometimes down the crest of the jaw. The teeth at the front of the mouth, by the beak, were smaller with larger teeth in the middle portion of the jaw. The lower teeth included one raised row of buccal teeth and several rows running diagonally on the medial and lingual portions. The teeth continued to grow throughout the animals life but, unlike with reptiles, the new teeth formed at the back of the tooth row. That happened within the groove on the surface of the lower jaw and three grooves on the maxilla. The lateral As they grew in, the teeth moved forward in the jaw through a combination of growth and remodeling. Instead of occluding their teeth together, ''Stenaulorhynchus'' had teeth that occluded against bone, which would wear the teeth down. Over time, the surfaces of the jaw bones would erode as well, causing the maxillary grooves to become shallower and more rounded.


Cranial

Some of the cranial features of ''Stenaulorhynchus'' include a frontal bone that is broader than it is long, the presence of a pineal foramen and a lack of ornamention on the jugal bone. Their postorbital The occipital condyles are significantly anterior to the quadrates and the quadrate and articular fit tightly together to form a jaw joint that wouldn't have allowed for much rotation. They area also known for their beak, formed at the front of their upper and lower jaws.


Postcranial

The dorsal vertebrae have a centrum that is taller than it is wide and slightly amphicoelous in shape. The parapophyses and prezygapophyses are distinct that have articular surfaces about 30 degrees above horizontal. The caudal vertebrae have tall neural spines and the second cervical rib has a backwards pointing spine. Their ischium fans out towards the rear of the animal. The humerus is robust. At the proximal end, the shaft is bent backwards and expanded along the coronal plane. The distal humerus has a flat posterior surface and a curved ectepicondylar groove. The radius and ulna are the same length. The femur is circular in cross-section and, as with the humerus, the proximal end is bent backwards. Both ends of the femur are expanded and the proximal end has no neck and a prominent greater trochanter. The distal end has thickened condyles with a groove on the lateral side.


Paleobiology

Microscopic analysis of thin-sections of bone have shown that ''Stenaulorhynchus'' had a determinant growth pattern, reaching 2/3 of their adult size within one year. The rate of bone deposition in the tibia and femur were 7-10 times higher within that year than in years 2-5. After 5 years of age, less than 1 micron of bone a day was deposited. It would have grown slower than
archosauriformes Archosauriformes (Greek for 'ruling lizards', and Latin for 'form') is a clade of diapsid reptiles that developed from archosauromorph ancestors some time in the Latest Permian (roughly 252 million years ago). It was defined by Jacques Gauthier ...
, the South American rhynchosaur species,
saurischia Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithis ...
n dinosaurs, and birds. ''Stenaulorhynchus'' continued to grow teeth throughout its life. Juveniles started out with three rows of upper teeth, with only one lateral to the main groove running along the upper surface of the maxilla. By the time they were adults, ''Stenaulorhynchus'' had several rows of maxillary teeth, one or more of which were lateral to the main maxillary groove. Older teeth would wear down with use and be resorbed where they contacted other teeth. ''Stenaulorhynchus'' may have been adapted for scratch-digging.


Discovery

The fossils were collected in 1930 by the British geologist G. M. Stockley during a survey commissioned by the Tanganyika Geological Survey Department. The goal was to better understand the “economic possibilities” of the Ruhaha Basin within Tanganyika Territory. He primarily collected fossils that had weathered out of the rock on the surface. The fossil collection was then described by Sidney Henry Haughton. While ''Stenaulorhynchus'' is the dominant vertebrate in the Manda beds, they also included a
Dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivorous animals with a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typic ...
and vertebrae from an unidentified
Theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
. The specimens weren’t all correctly labeled since bones from different localities fit together. More surprisingly, a
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is ...
was mislabeled as a maxillary fragment. Haughton also described a species which he named ''Stenaulorhynchus major,'' which he said differed primarily in size, based on a distal left and right humerus fragments. Those specimens were later identified as '' Stagonosuchus nyassicus'', a
suchia Suchia is a clade of archosaurs containing the majority of pseudosuchians (crocodilians and their extinct relatives). It was defined as the least inclusive clade containing '' Aetosaurus ferratus'', '' Rauisuchus tiradentes'', '' Prestosuchus c ...
n
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-avia ...
from the same locality and horizon.


Classification

''Stenaulorhynchus'' shares characteristics with other early rhynchosaurs, including their ankylothecodont dentition (teeth within deep sockets and fused to the bone) and precision-shear bite. They also have double-bladed dentaries and grooves on the upper surface of the maxilla. One of the differences between them is that ''Stenaulorhynchus'' had smaller maxillary teeth with a larger gap between the tooth rows. ''Stenaulorhynchus'' is differentiated from the later ''Hyperadapedon'' by its more lateral eyes, more forward braincase, and longer lower jaw with the teeth located more anteriorly. Hyperadapedons also lack the double-bladed dentary that the early genera have. The only absolute
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
between all the rhynchosaurs is that the dentary is at least half the total length of the jaw. Stenaulorhynchinae, a subfamily of rhynchosaurs including ''Stenaulorhhynchus'', has been proposed. Based on an analysis of morphological characteristics, one study defines it as including all taxa that are more closely related to ''Stenaulorhynchus stockleyi'' than ''Hyperodapedon gordoni''. The defining traits include having many teeth on the tongue side of the maxilla and a reduced size of the maxillary occlusal teeth allowing more teeth per row. ''Stenaulorhynchus'' has also been labeled as a sister-taxa to the Brazilian genus ''