Teratophoneus
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''Teratophoneus'' ("monstrous murderer"; Greek: ''teras'', "monster" and ''phoneus'', "murderer") is a
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
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to thirteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
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 ...
which lived during the late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
(late
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
age, about 77 to 76 million years ago) in what is now
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
, United States, containing a single known species, ''T. curriei''. It is known from an incomplete
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
and postcranial skeleton recovered from the
Kaiparowits Formation The Kaiparowits Formation is a sedimentary rock formation found in the Kaiparowits Plateau in Grand Staircase–Escalante National Monument, in the southern part of Utah in the western United States. It is over 2800 feet (850 m) thick, and is ...
. It was specifically named ''T. curriei'' in honor of Philip J. Currie.


Discovery and naming

Fossils of ''Teratophoneus'' were first found in the
Kaiparowits Formation The Kaiparowits Formation is a sedimentary rock formation found in the Kaiparowits Plateau in Grand Staircase–Escalante National Monument, in the southern part of Utah in the western United States. It is over 2800 feet (850 m) thick, and is ...
of southern Utah. Later, fossils from the same formation were discovered and identified as the genus. Argon-argon radiometric dating indicates that the Kaiparowits Formation was deposited between 76.1 and 74.0 million years ago, during the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
stage of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
period. This date means that ''Teratophoneus'' lived in the middle of the Campanian stage of the Late Cretaceous. Several fossils of ''Teratophoneus'' have been found. Originally, ''Teratophoneus'' was described based on the holotype BYU 8120. More recently, the specimens UMNH VP 16690 and UMNP VP 16691 have been assigned to it. In 2017, a new specimen of ''Teratophoneus'' was discovered in the Grand Staircase-Escalante National Monument and airlifted to the Natural History Museum of Utah in Salt Lake City. Later, in 2021, fossils belonging to 4 or 5 individuals were described in the same study. ''Teratophoneus'' was named by Thomas D. Carr, Thomas E. Williamson, Brooks B. Britt and Ken Stadtman in 2011. The type and only
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 ...
was named ''T. curriei''. The generic name is derived from the Greek ''teras'', "monster", and ''phoneus'', "murderer." The specific name honors Philip J. Currie.


Description

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
of ''Teratophoneus'' consists of a fragmentary skull and parts of the postcranial skeleton. The fossils were originally assigned to four different individuals, but are probably only of a single subadult animal. The specimen of ''Teratophoneus'' was not fully grown: according to an estimate by Carr ''et al''. was about in length and in weight. However this is likely an underestimate. In 2016,
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
gave an estimation of in length and in body mass for the maximum adult size. In the same year, Molina-Pérez and Larramendi estimated the size of the holotype at in length and in body mass. In 2021, based on the size of the frontal bone (similar to that of '' Lythronax''), Yun moderated the size of the subadult at approximately in length and in body mass. In the same year, the length of the only known articulated specimen UMNH VP 21100 was measured at and the maximum adult length of ''Teratophoneus'' was estimated at . Compared to the skull of an ''
Albertosaurus ''Albertosaurus'' (; meaning "Alberta lizard") is a genus of tyrannosaurid theropod dinosaurs that lived in western North America during the Late Cretaceous Period, about 71 million years ago. The type species, ''A. sarcophagus'', wa ...
'', ''Teratophoneus'' is roughly twenty-three percent shorter in proportion between the lacrimal bone of the and the tip of the snout. The skull of ''Teratophoneus'' is also comparably deeper. It is unclear if there was a specific reason for these differences, but the extra depth may have allowed for stronger jaw muscles, increasing the bite force of ''Teratophoneus''.


Classification

Loewen ''et al.'' (2013) conducted a phylogenetic analysis of the family Tyrannosauridae, and confirmed the assignment of ''Teratophoneus'' to the tyrannosaurid sub-family
Tyrannosaurinae The tyrannosaurines (Tyrannosaurinae) is an extinct subfamily of Tyrannosauridae a family of coelurosaurian theropod dinosaurs that consists of at least three tribes and several genera. All fossils of these genera have been found in the Late Cret ...
. They concluded that ''Teratophoneus'' was closely related to both ''
Tarbosaurus ''Tarbosaurus'' ( ; meaning "alarming lizard") is a genus of tyrannosaurid dinosaur that flourished in Asia about 70 million years ago, at the end of the Late Cretaceous Period, considered to contain a single known species, ''Tarbosaurus ba ...
'' and ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosaurus'' live ...
'', but placed in a more basal position within the family, though more derived than ''
Daspletosaurus ''Daspletosaurus'' ( ; meaning "frightful lizard") is a genus of tyrannosaurid dinosaur that lived in Laramidia between about 79.5 and 74 million years ago, during the Late Cretaceous Period. The genus ''Daspletosaurus'' contains three species ...
''. Below is the cladogram based on the
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
conducted by Loewen ''et al.'' in 2013. In 2020, when describing the genus ''
Thanatotheristes ''Thanatotheristes'' (meaning "reaper of death" according to the original authors; literally, "death harvester" in Greek language, Greek: , , "Death" and , , "harvester") is a genus of tyrannosaurid dinosaur from the Late Cretaceous of Larami ...
'', Voris ''et al.'', 2020 found ''Teratophoneus'' to be in a subclade alongside '' Dynamoterror'' and '' Lythronax''. The clade remains unnamed.


Paleobiology


Social Behavior

A bone bed of fossils from the Rainbows and Unicorns Quarry in Southern Utah's Kaiparowits Formation described in 2021 attributed to ''Teratophoneus'' suggests that the genus was a social pack-hunter. The fossils, consisting of four or possibly five animals ranging from 4-22 years of age, suggest a mass mortality event, possibly caused by flooding, or less likely by cyanobacterial toxicosis, fire or drought. The fact that all of the animals preserved died within a short time period further strengthens the argument for gregarious behavior in tyrannosaurids, with bone beds of ''Teratophoneus'', ''Albertosaurus'', and ''Daspletosaurus'' showcasing the potential behavior may have been widespread amongst tyrannosaurs in general.


Paleoecology

The holotype of ''Teratophoneus'' were recovered at the Kaiparowits Formation, in southern Utah. Argon-argon radiometric dating indicates that the fossils were buried during the Campanian stage of the Late Cretaceous period. During the Late Cretaceous period, the site within the Kaiparowits Formation was located on
Laramidia Laramidia was an island continent that existed during the Late Cretaceous period (99.6–66 Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island land mass separated from A ...
near its eastern shore on the
Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, and the Western Interior Sea) was a large inland sea that split the continent of North America into two landmasses. The ancient sea ...
, a large inland sea that split North America into two landmasses, the other being Appalachia to the east. The plateau where dinosaurs lived was an ancient floodplain dominated by large channels and an abundance of wetland
peat Peat (), also known as turf (), is an accumulation of partially decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, moors, or muskegs. The peatland ecosystem covers and is the most efficient ...
swamps, ponds and lakes, and was bordered by highlands. The climate was wet and humid, and supported an array of different and diverse groups of organisms. This formation contains one of the best and most continuous records of Late Cretaceous terrestrial life in the world. ''Teratophoneus curriei'' shared its paleoenvironment with
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 ...
s such as
dromaeosaurid Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
s, the
troodontid Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil disco ...
'' Talos sampsoni'', ornithomimids like '' Ornithomimus velox'', the ankylosaur Akainacephalus Johnsoni, the duckbilled hadrosaurs '' Parasaurolophus cyrtocristatus'' and ''
Gryposaurus monumentensis ''Gryposaurus'' (meaning "hooked-nosed (Greek ''grypos'') lizard"; sometimes incorrectly translated as " griffin (Latin ''gryphus'') lizard") was a genus of duckbilled dinosaur that lived about 80 to 75 million years ago, in the Late Cretac ...
'', the
ceratopsians 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 Jurassic ...
''
Utahceratops gettyi ''Utahceratops'' is an extinct genus of ceratopsian dinosaur that lived approximately 76.4~75.5 million years ago during the Late Cretaceous period in what is now Utah. ''Utahceratops'' was a large-sized, robustly-built, ground-dwelling, quadrupe ...
'', ''
Nasutoceratops titusi ''Nasutoceratops'' is an extinct genus of ceratopsian dinosaur. It is a basal Centrosaurinae, centrosaurine which lived during the Late Cretaceous period (geology), Period (late Campanian, about 76.0-75.5 Mya (unit), Ma). Fossils have been found ...
'', and ''
Kosmoceratops richardsoni ''Kosmoceratops'' () is a genus of ceratopsid dinosaur that lived in North America about 76–75.9 million years ago during the Late Cretaceous period. Specimens were discovered in Utah in the Kaiparowits Formation of the Grand Staircase–Esca ...
'', as well as the
oviraptorosauria Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or wi ...
n ''
Hagryphus giganteus ''Hagryphus'' (meaning " Ha's griffin") is a monospecific genus of caenagnathid dinosaur from southern Utah that lived during the Late Cretaceous (upper Campanian stage, 75.95 Ma) in what is now the Kaiparowits Formation of the Grand Staircase– ...
''. Paleofauna present in the Kaiparowits Formation included
chondrichthyans Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. C ...
(sharks and rays),
frog A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" ''Triadobatrachus'' is ...
s,
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
s,
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
s, lizards, and
crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest livi ...
ns. A variety of early mammals were present including
multituberculate Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, a ...
s,
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a ...
s, and
insectivora The order Insectivora (from Latin ''insectum'' "insect" and ''vorare'' "to eat") is a now-abandoned biological grouping within the class of mammals. Some species have now been moved out, leaving the remaining ones in the order Eulipotyphla, wi ...
ns.


See also

*
Timeline of tyrannosaur research This timeline of tyrannosaur research is a chronological listing of events in the history of paleontology focused on the tyrannosaurs, a group of predatory theropod dinosaurs that began as small, long-armed bird-like creatures with elaborate cr ...


References

{{Taxonbar, from=Q1705998 Tyrannosaurids Monotypic dinosaur genera Campanian life Late Cretaceous dinosaurs of North America Paleontology in Utah Kaiparowits Formation Fossil taxa described in 2011