Cryolophosaurus
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''Cryolophosaurus'' ( or ; "CRY-oh-loaf-oh-SAWR-us") 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 large
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 ...
dinosaur known from only a single
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 appropriat ...
''Cryolophosaurus ellioti'', from the early
Jurassic The Jurassic ( ) is a geologic period and 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 Jurassic constitutes the middle period of ...
of
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
. It was one of the largest theropods of the Early Jurassic, with the subadult being estimated to have reached long and weighed . ''Cryolophosaurus'' was first excavated from Antarctica's Early Jurassic,
Pliensbachian The Pliensbachian is an age (geology), age of the geologic timescale and stage (stratigraphy), stage in the stratigraphic column. It is part of the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series an ...
aged Hanson Formation, formerly the upper Falla Formation, by
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Dr. William Hammer in 1991. It was the first
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other ...
dinosaur to be discovered in Antarctica and the first non-avian dinosaur from the continent to be officially named. The sediments in which its fossils were found have been dated at ~194 to 188 million years ago, representing the Early Jurassic Period. ''Cryolophosaurus'' is known from a skull, a femur and other material, all of which have caused its classification to vary greatly. The femur possesses many primitive characteristics that have classified ''Cryolophosaurus'' as a dilophosaurid or a neotheropod outside of Dilophosauridae and Averostra, whereas the skull has many advanced features, leading the genus to be considered a tetanuran, an abelisaurid, a ceratosaur and even an allosaurid. ''Cryolophosaurus'' is currently considered to be a derived neotheropod, close to Averostra. Additionally, ''Cryolophosaurus'' possessed a distinctive " pompadour" crest that spanned the head from side to side. Based on evidence from related species and studies of bone texture, it is thought that this bizarre crest was used for intra-species recognition. The
brain A brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as Visual perception, vision. I ...
of ''Cryolophosaurus'' was also more primitive than those of other theropods.


Discovery and naming

''Cryolophosaurus'' originally was collected during the 1990–91 austral summer on
Mount Kirkpatrick Mount Kirkpatrick is a lofty, generally ice-free mountain in Antarctica's Queen Alexandra Range. Located 8 km (5 mi) west of Mount Dickerson, Mount Kirkpatrick is the highest point in the Queen Alexandra Range, as well as in its parent ...
in the
Beardmore Glacier The Beardmore Glacier in Antarctica is one of the largest valley glaciers in the world, being long and having a width of . It descends about from the Antarctic Plateau to the Ross Ice Shelf and is bordered by the Commonwealth Range of the Que ...
region of the Transantarctic Mountains. The discovery was made by Hammer, a professor at Augustana College, and his team. The fossils were found in the
siliceous Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, ...
of the Hanson Formation, formerly the upper Falla Formation, and dated to the
Pliensbachian Stage 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 Pli ...
of the early
Jurassic The Jurassic ( ) is a geologic period and 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 Jurassic constitutes the middle period of ...
. ''Cryolophosaurus'' was the second dinosaur, and first theropod, to be discovered in Antarctica. It was discovered after ''
Antarctopelta ''Antarctopelta'' ( ; meaning 'Antarctic shield') was a genus of ankylosaurian dinosaur with one known species, ''A. oliveroi'', which lived in Antarctica during the Late Cretaceous Period. It was a medium-sized ankylosaur, reaching 4 meters (13& ...
'', but named earlier. In 1991, both Hammer and the
Ohio State University The Ohio State University, commonly called Ohio State or OSU, is a public land-grant research university in Columbus, Ohio. A member of the University System of Ohio, it has been ranked by major institutional rankings among the best pub ...
geologist David Elliot excavated separate outcrops near Beardmore Glacier, sharing logistical expenses. Elliot's team first came across the remains of ''Cryolophosaurus'' in a rock formation around the altitude of high and about from the South Pole. When the discovery was made, they soon notified Hammer. Over the next three weeks, Hammer excavated of fossil-bearing rock. The team recovered over 100 fossil bones, including those of ''Cryolophosaurus''. The specimens were formally named and described in 1994 by Hammer and Hickerson, in the journal ''
Science Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and predictions about the universe. Science may be as old as the human species, and some of the earliest archeological evidence ...
''. During the 2003 season, a field team returned and collected more material from the original site. A second locality was discovered about higher in the section on Mt. Kirkpatrick. The name ''Cryolophosaurus ellioti'' is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words κρυος (meaning 'cold' or 'frozen', in reference to its discovery in Antarctica), λοφος (meaning 'crest') and (meaning 'lizard'), thus "cold crest lizard". Hammer and Hickerson named the species ''C. ellioti'', after David Elliot, who had made the initial discovery of the fossils.


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 seve ...
FMNH PR1821 is the only fully described specimen of ''Cryolophosaurus''. The specimen consists of 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 mandibles lacking most of their front half; nine maxillary
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, ...
; a fragmentary sixth cervical centrum; cervical
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 characteristi ...
7-10; several posterior cervical
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
s; several anterior dorsal vertebrae; most mid and posterior dorsal vertebrae; several dorsal ribs; the fifth sacral vertebrae; three
chevron Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * '' Eulithis testata'', a moth * Chevron (geology), a fold in rock la ...
s; many partial and complete caudal vertebrae and centra; two partial humeri; a proximal
radius In classical geometry, a radius (plural, : radii) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', ...
; a proximal
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
; a partial ilium; a proximal pubis; both
ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from Naples. It is the largest of the Phlegrean Islands. Roughly trapezoidal in shape, it measures approximately east to ...
, but only one
distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
; two incomplete femora; the distal end of a
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it conn ...
; the distal end of a
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity ...
, and the astragalus and calcaneum. In 2013, new material of ''Cryolophosaurus'' was unearthed in Antarctica. The description of this material has not yet been published in a non-abstract form. ''Cryolophosaurus'' was a large, well-built
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 ...
, one of the largest of its time. The holotype specimen is estimated to have reached long and weighed . Some researchers noted that the holotype individual probably represents a sub-adult, so adults could have been larger. Despite having slender proportions, ''Cryolophosaurus'' is one of the largest known
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 ...
theropods.


Skull

The holotype of ''Cryolophosaurus'' consists of a high, narrow skull, which was discovered articulated with the rest of the skeleton. The skull is an estimated long. It has a peculiar nasal crest that runs just over the eyes, where it rises perpendicular to the skull and fans out. It is thin and highly furrowed, giving it a unique "pompadour" appearance and earned it the nickname " Elvisaurus." The crest is an extension of the skull bones, near the tear ducts, fused on either side to orbital horns which rise from the eye sockets. While other theropods like the '' Monolophosaurus'' have crests, they usually run along the skull instead of across it. An unpublished study conducted by Vernon Meidlinger-Chin in 2013 suggested that previous studies lacked focus on endocranial details. The study found that the ''Cryolophosaurus'' fossil has a nearly complete, undistorted cranial cavity which is complete enough to give an approximate shape and size of the living brain. The endocast features clarified the dissimilarity of the skull with those of Allosauroids and Coelurosaurs giving ''Cryolophosaurus'' a basal position in Theropoda. Closer examination of how the skull bones fused reviewed details in the snout and forehead that are exceptionally similar to ''
Dilophosaurus ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 193 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserve ...
''.


Classification

Classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
of ''Cryolophosaurus'' is difficult because it has a mix of primitive and advanced characteristics. The femur has traits of early theropods, while the skull resembles much later species of the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
Tetanurae, like China's '' Sinraptor'' and '' Yangchuanosaurus''. This led
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at si ...
''et al.'' (1994) to place ''Cryolophosaurus'' in the taxon Allosauridae. Originally, Hammer and colleagues suspected that ''Cryolophosaurus'' might be a ceratosaur or even an early abelisaur, with some traits convergent with those of more advanced tetanurans, but ultimately concluded that it was itself the earliest known member of the tetanuran group. While a subsequent study by Hammer (along with Smith and Currie) again recovered ''Cryolophosaurus'' as a tetanuran, a later (2007) study by the same authors found that it was more closely related to ''
Dilophosaurus ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 193 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserve ...
'' and '' Dracovenator''. Sterling Nesbitt ''et al.'' (2009), using the characters of '' Tawa'' found ''Cryolophosaurus'' to be neither a dilophosaurid nor averostran neotheropod but instead the sister group of a clade composed of dilophosaurids and averostrans. However, in 2012, Matthew Carrano found that ''Cryolophosaurus'' was a tetanuran, related to '' Sinosaurus'', but unrelated to ''Dilophosaurus''. In 2020, a monograph of ''Dilophosaurus'' found ''Cryolophosaurus'' to be a derived neotheropod, close to Averostra, in a more derived position than ''
Zupaysaurus ''Zupaysaurus'' (; "ZOO-pay-SAWR-us") is an extinct genus of early theropod dinosaur living during the Norian stage of the Late Triassic in what is now Argentina. Fossils of the dinosaur were found in the Los Colorados Formation of the Ischigual ...
'', but less than ''Dilophosaurus''. The following cladogram illustrates a synthesis of the relationships of the early theropod groups compiled by Hendrickx ''et al.'' in 2015. However, a 2020, a study conducted Adam Marsh and Timothy Rowe found Cryolophosaurus to be a basal Neotheropod. While it was still closer to Averostra than Coelophysoidea was, it was still more basal than Dilophosaurus.


Paleobiology


Cranial ornamentation

Cranial display features, such as the one possessed by ''Cryolophosaurus'', make sense in social, gregarious animals, where other members of the species are available to observe and interpret messages of sexual status. Kevin Padian ''et al.'' (2004) challenged conventional hypotheses that the purpose of bizarre cranial structures and post-cranial armor in dinosaurs, was either for attracting mates, intimidating/fighting rivals in the group, or intimidating potential predators of other species. Padian ''et al.'' noted that based on phylogenetic, histological, and functional evidence these bizarre structures can be explained by the phenomenon of intra-species recognition, which is supported by the fossil evidence.
Thomas R. Holtz Jr. Thomas Richard Holtz Jr. (born September 13, 1965) is an American vertebrate palaeontologist, author, and principal lecturer at the University of Maryland's Department of Geology. He has published extensively on the phylogeny, morphology, ecomor ...
(2010) found that the bizarre crest of ''Cryolophosaurus'' was primarily for intra-species recognition, based on evidence from related species and studies of bone texture. According to Thomas Rich and his colleagues, the crest would have been ineffective as a weapon and may have possibly functioned as a display feature during certain types of social behavior such as mating. In 2019, a species recognition function was disputed but a socio-sexual display structure model was suggested.


Diet

When the type specimen was discovered, several long cervical ribs, of a supposed prosauropod dinosaur were found in the mouth of ''Cryolophosaurus'', which led Hammer (1998) to conclude that it was feeding on the prosauropod when it died. Hammer further noted that since the ribs were found extending all the way back to the theropod's neck region, this individual may have choked to death on these ribs. However, Smith ''et al.'' concluded that these remains belonged to the ''Cryolophosaurus'' specimen itself, and not to Hammer's "prosauropod". Hammer also concluded that a post-canine tooth belonging to a
tritylodont Tritylodontidae ("three-knob teeth", named after the shape of their cheek teeth) is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, bearing several mammalian traits like erect limbs, endothermy and details o ...
(an early mammal relative), found with the remains, was part of its stomach contents when it died.


Paleopathology

Some ''Cryolophosaurus'' bones have pathologies that show evidence of scavenging. Broken teeth from a juvenile ''Cryolophosaurus'' were found nearby. These teeth have no roots and likely shed naturally while scavenging the adult ''Cryolophosaurus'' carcass. Another possible pathology is found in the astragalus (ankle bone) of ''Cryolophosaurus''. This bone was preserved with a small splint from the
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity ...
located just above the ankle. The splint, however, may also be just a unique morphological feature of ''Cryolophosaurus''.


Paleoenvironment

All known specimens of ''Cryolophosaurus'' have been recovered in the Hanson Formation, which is one of only two major dinosaur-bearing rock formations found on the continent of Antarctica. It was discovered in "tuffaceous" siltstone deposited in the Sinemurian to Pliensbachian stage of the Early Jurassic, approximately 194 to 188 million years ago. This
geological formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics ( lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock exp ...
is part of the Victoria Group of the Transantarctic Mountains, which is approximately above sea level. The high altitude of this site supports the idea that early Jurassic Antarctica had
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
s populated by a diverse range of species, at least along the coast. The Hanson Formation was deposited in an active volcano−tectonic rift system formed during the breakup of
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final sta ...
. In the Early Jurassic, Antarctica was closer to the
equator The equator is a circle of latitude, about in circumference, that divides Earth into the Northern and Southern hemispheres. It is an imaginary line located at 0 degrees latitude, halfway between the North and South poles. The term can also ...
and the world was considerably warmer than today, but the
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
was still cool
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
. Models of Jurassic air flow indicate that coastal areas probably never dropped much below freezing, although more extreme conditions existed inland. ''Cryolophosaurus'' was found about from the
South Pole The South Pole, also known as the Geographic South Pole, Terrestrial South Pole or 90th Parallel South, is one of the two points where Earth's axis of rotation intersects its surface. It is the southernmost point on Earth and lies antipod ...
but, at the time it lived, this was about or so farther north. This formation has produced the remains of ''
Glacialisaurus ''Glacialisaurus'' is a genus of sauropodomorph dinosaur. It lived during the Pliensbachian stage of the Early Jurassic period around 186 to 182 million years ago in what is now the central region of the Transantarctic Mountains of Antarctica. I ...
'' (a large basal sauropodomorph), a crow-sized
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
(a dimorphodontid), a
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
(a
tritylodon ''Tritylodon'' (Greek for 3 cusped tooth) is an extinct genus of tritylodonts, one of the most advanced group of cynodont therapsids. They lived in the Early Jurassic and possibly Late Triassic periods along with dinosaurs. They also shared man ...
t, which is a type of
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
about the size of a rat), herbivorous synapsid, and two small unnamed sauropodomorphs. There are also the remains of many plant genera recovered from the Early Jurassic Camp Hill Formation, around the same age as fossils of ''Cryolophosaurus'', proving that dense plant matter had once grown on Antarctica's surface before it drifted southward.


References


External links


Transantarctic Vertebrate Paleontology Project
official website for the NSF project conducting research on Cryolophosaurus and related fauna (photos, research information, publication list, geology, project members ...)

(photos, information)
Field Museum of Natural History
(video, information) {{Portal bar, Dinosaurs Prehistoric tetanurans Pliensbachian life Sinemurian life Dinosaurs of Antarctica Early Jurassic dinosaurs Jurassic Antarctica Fossil taxa described in 1994 Taxa named by William R. Hammer Fossils of Antarctica