Coelurosauria


 * For the Permian gliding reptile, see Coelurosauravus.

Coelurosauria (from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. In the past, it was used to refer to all small theropods, although this classification has been abolished. Nonetheless, it is still a diverse group that includes compsognathids, tyrannosaurs, ornithomimosaurs, and maniraptorans; Maniraptora includes birds, the only dinosaurs alive today. Most feathered dinosaurs discovered so far have been coelurosaurs; Philip J. Currie considers it probable that all coelurosaurs were feathered.

Bodyplan
All coelurosaurs were bipedal, and most were carnivores, though many groups exhibited a more varied diet including insectivory (alvarezsauridae), omnivory (oviraptoridae and troodontidae), and herbivory (therizinosauridae). The group includes some of the largest (Tyrannosaurus) and smallest (Microraptor, Parvicursor) carnivorous dinosaurs ever discovered. Characteristics that distinguish coelurosaurs include:
 * a sacrum (series of vertebrae that attach to the hips) longer than in other dinosaurs
 * a tail stiffened towards the tip
 * a bowed ulna (lower arm bone).
 * a tibia (lower leg bone) that is longer than the femur (upper leg bone)

Integument
To date, fossilized traces of feathers have been identified among most coelurosaurian lineages. Feathers of some type, or morphological features suggesting feathers, have been found in fossils of at least one species in all coelurosaur subgroups other than ornithomimosauria. Modern birds are classified as coelurosaurs by nearly all palaeontologists. The arrangements of feathers currently observed on Coelurosauria other than modern birds are without exception more primitive, and some coelurosaurian species are known to have had bare or scaly skin rather than feathers on at least some parts of their bodies. These include large tyrannosaurids, some compsognathids such as Juravenator, and Scansoriopteryx. All preserve the impression of scales from regions near the hind legs and tail, though Scansoriopteryx and possibly Juravenator also preserve feathers elsewhere on the body. Most coelurosaurs, including modern birds, retained scales on the feet, though in a few (such as Anchiornis and the modern Rock Ptarmigan) the feet and toes are also entirely covered in feathers.

Though once thought to be a feature exclusive to coelurosaurs, feathers or feather-like structures are also known in some ornithischian dinosaurs (like Tianyulong), though whether these are related to true feathers or novel structures that evolved independently is unknown.

Nervous system and senses
Although rare, complete casts of theropod endocrania are known from fossils. Theropod endocrania can also be reconstructed from preserved braincases without damaging valuable specimens by using a computed tomography scan and 3D reconstruction software. These finds are of evolutionary significance because they help document the emergence of the neurology of modern birds from that of earlier reptiles. An increase in the proportion of the brain occupied by the cerebrum seems to have occurred with the advent of the Coelurosauria and "continued throughout the evolution of maniraptorans and early birds."

Fossil evidence and age
A few fossil traces tentatively associated with the Coelurosauria date back as far as the late Triassic. What has been found between then and the start of the late Jurassic is fragmentary. A typical example is Iliosuchus, known only from two ilia bones in the mid Jurassic. It was a 1.5 m long carnivore from about 165 Ma (million years ago) in Oxfordshire and is tentatively assigned to the Tyrannosauroidea.

Many nearly complete fossil coelurosaurians are known from the late Jurassic. Archaeopteryx (incl. Wellnhoferia) is known from Bavaria at 155-150 Ma. Ornitholestes, the troodontid WDC DML 110, Coelurus fragilis and Tanycolagreus topwilsoni are all known from the Morrison Formation in Wyoming at about 150 Ma. Epidendrosaurus and Pedopenna are known from the Daohugou Beds in China, whose age is still being debated, but may be about 160 Ma or 145 Ma.

The wide range of fossils in the late Jurassic and morphological evidence suggests that coelurosaurian differentiation was virtually complete before the end of the Jurassic.

In the early Cretaceous, a superb range of coelurosaurian fossils (including avians) are known from the Yixian Formation in Liaoning. All known theropod dinosaurs from the Yixian Formation are coelurosaurs. Many of the coelurosaurian lineages survived to the end of the Cretaceous period (about 65 Ma) and fossils of some lineages, such as the Tyrannosauroidea, are best known from the late Cretaceous. A majority of coelurosaur groups became extinct in the Cretaceous–Paleogene extinction event, including the Tyrannosauroidea, Ornithomimosauria, Oviraptorosauria, Deinonychosauria, Enantiornithes, and Hesperornithes. Only the Neornithes (modern birds) survived, and continued to diversify after the extinction of the other dinosaurs into the numerous forms found today.

There is consensus among paleontologists that birds are the descendants of coelurosaurs. Under modern cladistic definitions, birds are considered the only living lineage of coelurosaurs. Birds are classified by most paleontologists as belonging to the subgroup Maniraptora.

Classification
The phylogeny and taxonomy of Coelurosauria has been subject to intensive research and revision. For many years, Coelurosauria was a 'dumping ground' for all small theropods. In the 1960s several distinctive lineages of coelurosaurs were recognized, and a number of new infraorders were erected, including the Ornithomimosauria, Deinonychosauria, and Oviraptorosauria. During the 1980s and 1990s, paleontologists began to give Coelurosauria a formal definition, usually as all animals closer to birds than to Allosaurus, or equivalent specifiers. Under this modern definition, many small theropods are not classified as coelurosaurs at all and some large theropods, such as the tyrannosaurids, were actually more advanced than allosaurs and therefore were reclassified as giant coelurosaurs. Even more drastically, the segnosaurs, once not even regarded as theropods, have turned out to be non-carnivorous coelurosaurs related to Therizinosaurus. Senter (2007) listed 59 different published phylogenies since 1984. Those since 2005 have followed almost the same pattern, and differ significantly from many older phylogenies.

The following phylogenetic results are taken from Senter (2007) "A new look at the Phylogeny of Coelurosauria (Dinosauria: Theropoda)." This cladogram does not represent time, but a crude estimate of the time can be inferred from morphological changes. The first coelurosaurs were similar to the coelurids Tanycolagreus and Coelurus, and differed only sightly from other early tyrannosauroids Dilong and Eotyrannus. The two most significant separations between subgroups are those between the Paraves and other coelurosaurs and between the paravian clades Avialae and Deinonychosauria.

1 Coelurosauria, 2 †Tyrannosauroidea, 3 †Coeluridae, 4 †Compsognathidae, 5 Maniraptoriformes, 6 †Ornithomimosauria, 7 Maniraptora, 8 †Therizinosauroidea, 9 †Therizinosauridae, 10 †Alvarezsauridae, 11 †Oviraptorosauria, 12 †Oviraptoridae, 13 †Ingeniinae, 14 Paraves, 15 Avialae, 16 Aves, 17 Ornithurae, 18 †Deinonychosauria, 19 †Troodontidae, 20 †Dromaeosauridae, 21 †Unenlagiinae, 22 †Microraptoria, 23 †Dromaeosaurinae

"Coelurosaurus"
"Coelurosaurus" is an informal generic name, attributed to Friedrich von Huene, 1929, that is sometimes seen in lists of dinosaurs. The name is undescribed and has not been used seriously, although it has appeared in works of fiction.