''Triopticus'' is a genus of
archosauriform
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 Gauthie ...
reptile from the
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 ...
of
Texas
Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
,
United States
The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., federal district, five ma ...
. It contains a single species, ''Triopticus primus'', described in 2016 by Stocker ''et al''. It has an unusually domed head reminiscent of the later
pachycephalosauria
Pachycephalosauria (; from Greek παχυκεφαλόσαυρος for 'thick headed lizards') is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachycep ...
n
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 ...
s in an example 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 ...
.
Description
The portion of the head that is preserved shows remarkable similarities to the specialized heads of
pachycephalosaurs
Pachycephalosauria (; from Greek παχυκεφαλόσαυρος for 'thick headed lizards') is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachycep ...
. Five bosses, or rounded protuberances of bone are visible on the fossil, with one on the
frontal bone
The frontal bone is a bone in the human skull. The bone consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, pa ...
and two pairs behind (respectively on the
postorbital bone
The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some v ...
and the
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone.
In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral co ...
/
parietal bone
The parietal bones () are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is named ...
s) separated by a shallow groove. The backmost pair of bosses form a thick shelf that stretches outwards and backwards over the rear end of the skull, comparable to the domes of pachycephalosaurs and the frills of
ceratopsia
Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassi ...
ns.
The bosses have very roughly textured surfaces, suggesting that they would have been covered by keratin in life.
CT scan
A computed tomography scan (CT scan; formerly called computed axial tomography scan or CAT scan) is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers ...
s show that the bosses internally have three distinct layers of bone, with a dense innermost layer which resembles the typical skull roof in texture, followed by a spongy, porous layer filled with blood vessels, and the least dense third layer close to the surface. Pachycephalosaurs like ''
Stegoceras
''Stegoceras'' is a genus of pachycephalosaurid (dome-headed) dinosaur that lived in what is now North America during the Late Cretaceous period, about 77.5 to 74 million years ago (mya). The first specimens from Alberta, Canada, were descri ...
'' also similarly have three layers of bone inside its dome,
but the second layer is the least dense as opposed to the third. However, this difference may arise from differential deposition of minerals in the skull.
The deep pit in the back of the skull of ''Triopticus'' likely arises from the two back pairs of bosses growing around this region, not unlike the holes of
doughnut
A doughnut or donut () is a type of food made from leavened fried dough. It is popular in many countries and is prepared in various forms as a sweet snack that can be homemade or purchased in bakeries, supermarkets, food stalls, and fran ...
s. It is also possible that the pit is an opening for the
pineal gland
The pineal gland, conarium, or epiphysis cerebri, is a small endocrine gland in the brain of most vertebrates. The pineal gland produces melatonin, a serotonin-derived hormone which modulates sleep patterns in both circadian and seasonal ...
, as it is in approximately the right position, but this is less likely because no other archosauriforms have such an opening,
and the opening does not appear to be connected to any internal structures. The pit has the same texture as the bosses, suggesting that it was also potentially covered in keratin.
The eye socket is very large, and it is surrounded by
mineralized cartilage, a trait only seen otherwise in pachycephalosaurs. CT scans also revealed large
optic nerve
In neuroanatomy, the optic nerve, also known as the second cranial nerve, cranial nerve II, or simply CN II, is a paired cranial nerve that transmits visual information from the retina to the brain. In humans, the optic nerve is derived fro ...
s as well as long
semicircular canals
The semicircular canals or semicircular ducts are three semicircular, interconnected tubes located in the innermost part of each ear, the inner ear. The three canals are the horizontal, superior and posterior semicircular canals.
Structure
Th ...
and a relatively large
cerebellar flocculus (both involved in stabilizing gaze,
although long semicircular canals are also associated with bipedal stances
). These traits suggest that ''Triopticus'' was a very visually-oriented animal. Otherwise, the structure of the brain is generally similar to other archosauriforms, although it superficially resembles that of ''Stegoceras''.
Despite the heavy similarity of ''Triopticus'' to pachycephalosaurs, there are a number of traits that indicate its archosauriform identity. In ''Triopticus'', the
parabasisphenoid bone is horizontal, the posttemporal fenestrae at the back of the braincase are not sealed off at all, and the region where the
squamosal bone The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone.
In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral co ...
and
quadrate bone
The quadrate bone is a skull bone in most tetrapods, including amphibians, sauropsids (reptiles, birds), and early synapsids.
In most tetrapods, the quadrate bone connects to the quadratojugal and squamosal bones in the skull, and forms uppe ...
come into contact is not externally visible, all of which indicate that it is not a basal dinosaur.
Additionally, a narrow strip of smooth bone on the
lacrimal bone
The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...
in front of the eye sockets forms the margin of the
antorbital fenestra
An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, birds ...
and the associated antorbital fossa, which are distinctly archosauriform characteristics.
Discovery and naming
The only known specimen of ''Triopticus'',
TMM 31100-1030, consists of a
braincase
In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
containing part of the
eye socket
In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...
. It was discovered in Quarry 3 of the Otis Chalk in
Howard County, Texas
Howard County is a county located in the U.S. state of Texas. At the 2020 census, its population was 34,860. Its county seat is Big Spring. The county was created in 1876 and organized in 1882. It is named for Volney E. Howard, a U.S. Congr ...
, which is part of the
Dockum Group
The Dockum is a Late Triassic (approximately late Carnian through Rhaetian, or 223–200 Ma) geologic group found primarily on the Llano Estacado of western Texas and eastern New Mexico with minor exposures in southwestern Kansas, eastern Color ...
. While the Otis Chalk itself has not yet been dated, comparisons with the animal fauna of other sites (including the
Pekin Formation of
North Carolina
North Carolina () is a state in the Southeastern region of the United States. The state is the 28th largest and 9th-most populous of the United States. It is bordered by Virginia to the north, the Atlantic Ocean to the east, Georgia a ...
) suggest a potential age between 229 Ma and 226 Ma.
The genus name of ''Triopticus'' is derived from the Latin ''tri'' ("three") and ''optic'' ("vision"), in reference to a large pit in the back of the skull reminiscent of a third eye socket. The species name, ''primus'', is the Latin word for "first".
Classification
A
phylogenetic analysis
In biology
Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that pro ...
was conducted in 2016 in an attempt to elucidate the relationships of ''Triopticus''. The
phylogenetic tree recovered by the analysis, reproduced below, found ''Triopticus'' in a
polytomy
An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with other
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 ...
archosauriforms.
Paleoecology
In the Otis Chalk assemblage, ''Triopticus'' lived alongside a number of other archosauriforms including the
allokotosaurs ''
Trilophosaurus'' and ''
Malerisaurus
''Malerisaurus'' is an extinct genus of archosauromorph known from Andhra Pradesh of India and Texas of the USA.
Description
''Malerisaurus'' was a medium-sized archosauromorph which averaged 1.2 meters in length. ''Malerisaurus'' is known fr ...
''; the
doswelliid ''
Doswellia
''Doswellia'' is an extinct genus of archosauriform from the Late Triassic of North America. It is the most notable member of the family Doswelliidae, related to the proterochampsids. ''Doswellia'' was a low and heavily built carnivore which li ...
''; the
phytosaur
Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalent gr ...
s ''
Parasuchus
''Parasuchus'' is an extinct genus of basal phytosaur known from the Late Triassic (late Carnian to early Norian stage) of Andhra Pradesh and Madhya Pradesh, India. At its most restricted definition, ''Parasuchus'' contains a single species, ' ...
'' and ''
Angistorhinus''; the
aetosaur
Aetosaurs () are heavily armored reptiles belonging to the extinct order Aetosauria (; from Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs ...
s ''
Longosuchus'', ''
Lucasuchus'', and ''
Coahomasuchus''; the
shuvosaurid
Shuvosauridae is an extinct family of theropod-like pseudosuchians within the clade Poposauroidea. Shuvosaurids existed in North America (United States) and South America (Argentina) during the Late Triassic period (late Carnian to Rhaetian sta ...
''
Effigia
''Effigia'' was an extinct genus of shuvosaurid known from the Late Triassic of New Mexico, south-western USA. With a bipedal stance, long neck, and a toothless beaked skull, ''Effigia'' and other shuvosaurids bore a resemblance to the ornithomi ...
''; the
silesaurid
Silesauridae is an extinct family of Triassic dinosauriforms. It is most commonly considered to be a clade of non-dinosaur dinosauriforms, and the sister group of dinosaurs. Some studies have instead suggested that most or all silesaurids compr ...
''
Silesaurus
''Silesaurus'' is a genus of silesaurid dinosauriform from the Late Triassic, of what is now Poland.
Discovery
Fossilized remains of ''Silesaurus'' have been found in the Keuper Claystone in Krasiejów near Opole, Silesia, Poland, which is ...
''; the
lagerpetid dinosauromorph
Dinosauromorpha is a clade of avemetatarsalian archosaurs (reptiles closer to birds than to crocodilians) that includes the Dinosauria (dinosaurs) and some of their close relatives. It was originally defined to include dinosauriforms and lage ...
''
Dromomeron
''Dromomeron'' (meaning "running femur") is a genus of lagerpetid avemetatarsalian which lived around 220 to 211.9 ± 0.7 million years ago. The genus contains species known from Late Triassic-age rocks of the Southwestern United States and north ...
'', and the
coelophysoid 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 ...
dinosaur ''
Lepidus
Marcus Aemilius Lepidus (; c. 89 BC – late 13 or early 12 BC) was a Roman general and statesman who formed the Second Triumvirate alongside Octavian and Mark Antony during the final years of the Roman Republic. Lepidus had previously bee ...
''. Some of these also exhibit evolutionary convergence upon archosaurs living after the
Triassic–Jurassic extinction event
The Triassic–Jurassic (Tr-J) extinction event, often called the end-Triassic extinction, marks the boundary between the Triassic and Jurassic periods, , and is one of the top five major extinction events of the Phanerozoic eon, profoundly affect ...
- the phytosaurs resemble the
thalattosuchians
Thalattosuchia is a clade of marine crocodylomorphs from the Early Jurassic to the Early Cretaceous that had a cosmopolitan distribution. They are colloquially referred to as marine crocodiles or sea crocodiles, though they are not members of Cro ...
,
gavialoids, and
spinosaurids
The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includ ...
, the aetosaurs resemble the
ankylosaurs, and the shuvosaurids resemble the
ornithomimosaurs
Ornithomimosauria ("bird-mimic lizards") are theropod dinosaurs which bore a superficial resemblance to the modern-day ostrich. They were fast, omnivorous or herbivorous dinosaurs from the Cretaceous Period of Laurasia (now Asia, Europe and North ...
.
References
External links
An annotated 3D model of the ''Triopticus'' skull.A transparent 3D model of the ''Triopticus'' skull, showing its brain structure.
{{Taxonbar, from=Q28432240
Prehistoric archosauriforms
Prehistoric reptile genera
Late Triassic reptiles of North America
Paleontology in Texas
Fossil taxa described in 2016