Dilong Paradoxus
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''Dilong'' (帝龍, which means 'emperor dragon') is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of small basal
tyrannosauroid 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 supercontinen ...
dinosaurs. The only species in the genus is ''Dilong paradoxus'', known from the
Lower Lower may refer to: * ''Lower'' (album), 2025 album by Benjamin Booker *Lower (surname) *Lower Township, New Jersey *Lower Receiver (firearms) *Lower Wick Lower Wick is a small hamlet located in the county of Gloucestershire, England. It is sit ...
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
Yixian Formation The Yixian Formation (; formerly Romanization of Chinese, transcribed as Yihsien Formation or Yixiang Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. I ...
of
East Asia East Asia is a geocultural region of Asia. It includes China, Japan, Mongolia, North Korea, South Korea, and Taiwan, plus two special administrative regions of China, Hong Kong and Macau. The economies of Economy of China, China, Economy of Ja ...
(in what is now
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
). The discovery of ''Dilong'' was significant as it provided the first evidence of feathers in tyrannosaurs, something which had previously been suspected for the group.


Discovery

The ''Dilong''
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
specimen was found in the Lujiatun Unit of the Yixian Formation near Lujiatun, Beipiao, in western
Liaoning ) , image_skyline = , image_alt = , image_caption = Clockwise: Mukden Palace in Shenyang, Xinghai Square in Dalian, Dalian coast, Yalu River at Dandong , image_map = Liaoning in China (+all claims hatched).svg , ...
Province,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. This formation contains rocks dating to between 125.8–126 million years old. However, specimens have been found in other areas of the Yixian Formation which may vary in age. ''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 theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
rex'', the "king" tyrannosaurid. "Long" is used to name Chinese dinosaurs in much the same way that the ancient Greek ''-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 spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
isation of the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
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 People's Republic of China, China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian For ...
in
Beijing Beijing, Chinese postal romanization, previously romanized as Peking, is the capital city of China. With more than 22 million residents, it is the world's List of national capitals by population, most populous national capital city as well as ...
), 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 an exa ...
s or protofeathers. The feathers were seen in fossilized skin impressions from near the
jaw The jaws are a pair of opposable articulated structures at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth ...
and tail. They are not identical to modern
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
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 a Provinces and territories of Canada, province in Canada. It is a part of Western Canada and is one of the three Canadian Prairies, prairie provinces. Alberta is bordered by British Columbia to its west, Saskatchewan to its east, t ...
and
Mongolia Mongolia is a landlocked country in East Asia, bordered by Russia to the north and China to the south and southeast. It covers an area of , with a population of 3.5 million, making it the world's List of countries and dependencies by po ...
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 supercontinent ...
, the group that includes the later, larger tyrannosaurids such as ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
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 ''Coelurus'' ( ) is a genus of coelurosaurian dinosaur from the Late Jurassic period (mid-late Kimmeridgian faunal stage, 155–152 million years ago). The name means "hollow tail", referring to its hollow tail vertebrae (Greek κοῖλ ...
'', 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 Period (geology), Periods. The bird-like features of these species, a ...
.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. However, in an analysis by Brusatte ''et al.'' in 2016, both parsimony and Bayesian phylogenetic analyses placed ''Dilong'' outside of Proceratosauridae, as a slightly more advanced tyrannosauroid. In 2024, the describers of '' Alpkarakush'' recovered ''Dilong'' and proceratosaurids (''Proceratosaurus'' and ''Guanlong'') outside Tyrannosauroidea based on their phylogenetic analysis.


Paleobiology

Braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, 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 calv ...
scans indicate that ''Dilong'' had an S-shaped brain protected by thin
meninges In anatomy, the meninges (; meninx ; ) 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 the subarachnoid spac ...
, 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 () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns. The large flocculus of ''Dilong'' suggests it was agile and had good balance, while small
olfactory tract The olfactory tract (olfactory peduncle or olfactory stalk) 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 cort ...
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 Tyrannosauroidea Dinosaur species Barremian dinosaurs Yixian Formation Taxa named by Xu Xing Taxa named by Mark Norell Fossil taxa described in 2004 Dinosaurs of China Feathered dinosaurs