Dilong paradoxus
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''Dilong'' (帝龍, which means 'emperor dragon') 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 basal tyrannosauroid
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 ...
. The only species is ''Dilong paradoxus''. It is from the Lower
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 ...
Yixian Formation The Yixian Formation (; formerly transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its exq ...
near Lujiatun, Beipiao, in the western
Liaoning Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmo ...
province of
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
. It lived about 126 million years ago.


Discovery

''Dilong'' was described by Xu Xing and colleagues in 2004. The name is derived from the Chinese 帝 ''dì'' meaning 'emperor' and 龙 / 龍 ''lóng'' meaning 'dragon'. "Di", "emperor", refers to the relationship of this animal to ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropoda, 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. ''Tyrannosa ...
rex'', the "king" tyrannosaurid. "Long" is used to name Chinese dinosaurs in much the same way that the Latin ''-saur'' is in the West. The specific name, ''paradoxus'', is a
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
isation of the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...
παράδοξον meaning 'against received wisdom'.


Description

The type specimen is IVPP 14243 (
Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name sugg ...
in
Beijing } Beijing ( ; ; ), Chinese postal romanization, alternatively romanized as Peking ( ), is the Capital city, capital of the China, People's Republic of China. It is the center of power and development of the country. Beijing is the world's Li ...
), a nearly complete, semi-articulated, skull and skeleton. Referred material includes IVPP 1242, a nearly complete skull and presacral vertebrae, TNP01109 ( Tianjin Museum of Natural History), a partial skull, and IVPP V11579, another skull which may belong to ''D. paradoxus'', or to a related species. The type specimen of ''Dilong'' was about in length, but it is thought to be a juvenile and may have been over long when fully grown. Gregory S. Paul estimated that a long individual would have weighed .


Feathers

''Dilong paradoxus'' had a covering of simple
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premie ...
s or
protofeathers A feathered dinosaur is any species of dinosaur possessing feathers. While this includes all species of birds, there is a hypothesis that many, if not all non-avian dinosaur species also possessed feathers in some shape or form. It has been su ...
. The feathers were seen in fossilized skin impressions from near the jaw and tail. They are not identical to modern
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
feathers, lacking a central shaft and most likely used for warmth, since they could not have enabled flight. Adult tyrannosaurs, found in
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest T ...
and
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million ...
have skin impressions which appear to show the pebbly scales typical of other dinosaurs. Xu et al. (2004) speculated that the tyrannosauroids may have had different skin coverings on different parts of their bodies—perhaps mixing scales and feathers. They also speculated that feathers may correlate negatively with body size—that juveniles may have been feathered, then shed the feathers and expressed only scales as the animal became larger and no longer needed insulation to stay warm.


Classification

When ''Dilong'' was first described, it was considered one of the earliest and most primitive members of
Tyrannosauroidea Tyrannosauroidea (meaning 'tyrant lizard forms') is a superfamily (or clade) of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontin ...
, the group that includes the later, larger
tyrannosaurids 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 ...
such as ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropoda, 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. ''Tyrannosa ...
rex''. At least one later study, by Turner and colleagues in 2007, reanalyzed the relationships of coelurosaurian dinosaurs, including ''Dilong'', and found that it was not a tyrannosauroid. Rather, they placed ''Dilong'' two steps above the tyrannosauroids in their phylogeny; more advanced than '' Coelurus'', but more primitive than the
Compsognathidae Compsognathidae is a family of coelurosaurian theropod dinosaurs. Compsognathids were small carnivores, generally conservative in form, hailing from the Jurassic and Cretaceous Periods. The bird-like features of these species, along with other d ...
.Turner, A.H., Pol, D., Clarke, J.A., Erickson, G.M., and Norell, M. (2007). "Supporting online material for: A basal dromaeosaurid and size evolution preceding avian flight". ''Science'', 317: 1378-1381.
supplement
However, other studies continued to find ''Dilong'' as a tyrannosauroid, and some (such as Carr & Williamson 2010) found ''Dilong'' to fall within Tyrannosauroidea, not among the more advanced coelurosaurs. Below is a cladogram containing most tyrannosauroids by Loewen ''et al.'' in 2013. In a 2014 study, ''Dilong'' was found to be a
proceratosaurid Proceratosauridae is a family or clade of tyrannosauroid theropod dinosaurs from the Middle Jurassic to the Early Cretaceous. Distinguishing features Unlike the advanced tyrannosaurids but similar to primitive tyrannosauroids like '' Dilong' ...
. However, in an analysis by Brusatte ''et al.'' in 2016, both
parsimony Parsimony refers to the quality of economy or frugality in the use of resources. Parsimony may also refer to * The Law of Parsimony, or Occam's razor, a problem-solving principle ** Maximum parsimony (phylogenetics), an optimality criterion in p ...
and
Bayesian Thomas Bayes (/beɪz/; c. 1701 – 1761) was an English statistician, philosopher, and Presbyterian minister. Bayesian () refers either to a range of concepts and approaches that relate to statistical methods based on Bayes' theorem, or a followe ...
phylogenetic analyses placed ''Dilong'' outside of Proceratosauridae, as a slightly more advanced tyrannosauroid.


Paleobiology

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 ...
scans indicate that ''Dilong'' had an S-shaped brain protected by thin
meninges In anatomy, the meninges (, ''singular:'' meninx ( or ), ) are the three membranes that envelop the brain and spinal cord. In mammals, the meninges are the dura mater, the arachnoid mater, and the pia mater. Cerebrospinal fluid is located in ...
, unlike ''Tyrannosaurus'' which has a more linear brain protected by thicker meninges; this is probably a size-related trait, as it is in
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. The large
flocculus The flocculus (Latin: ''tuft of wool'', diminutive) is a small lobe of the cerebellum at the posterior border of the middle cerebellar peduncle anterior to the biventer lobule. Like other parts of the cerebellum, the flocculus is involved in moto ...
of ''Dilong'' suggests it was agile and had good balance, while small
olfactory tract The olfactory tract is a bilateral bundle of afferent nerve fibers from the mitral and tufted cells of the olfactory bulb that connects to several target regions in the brain, including the piriform cortex, amygdala, and entorhinal cortex. It ...
s suggest that its sense of smell was not as refined as that of ''Tyrannosaurus'' and other more advanced tyrannosauroids.


See also

* Timeline of tyrannosaur research


References


External links


Original ''Nature'' article as a PDF

''New Scientist'' news article


{{Taxonbar, from=Q27439 Early Cretaceous dinosaurs of Asia Feathered dinosaurs Fossil taxa described in 2004 Tyrannosaurs Taxa named by Xu Xing Taxa named by Mark Norell Yixian fauna